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VOL. 22 

ISBN 0 621 02870 3 


which deals with the territories of 



Edited by 
J. H. Ross 

Editorial Committee: B. de Winter, D. J. B. Killick, 
O. A. Leistner and J. H. Ross 

Botanical Research Institute, 
Department of Agricultural Technical Services 






Introduction vii 

Plan of Flora viii 

Ochnaceae by P. C. V. du Toit and A. A. Obermeyer 1 

Clusiaceae by D. J. B. Killick and N. K. B. Robson 14 

Elatinaceae by A. A. Obermeyer 23 

Frankeniaceae by A. A. Obermeyer 32 

Tamaricaceae by A. A. Obermeyer 36 

Canellaceae by L. E. Codd 39 

Violaceae by A. A. Obermeyer 42 

Flacourtiaceae by D. J. B. Killick and J. E. Langenegger (Dovyalis) 53 

Tumeraceae by A. A. Obermeyer 93 

Passifloraceae by W. J. J. O. de Wilde 104 

Achariaceae by D. J. B. Killick 128 

Loasaceae by L. E. Codd 134 

Begoniaceae by O. M. Hilliard 136 

Cactaceae by A. A. Obermeyer 144 

Index 157 


Digitized by the Internet Archive 
in 2016 


For a key to the families, the Flora should be used in conjunction with Phillips’s Genera of 
South African Flowering Plants, ed. 2 (1951) and Dyer’s Genera of Southern African Flowering 
Plants, Vol. 1 (1975) and Vol. 2 (in press), which are arranged on the lines of the Engler system. 
The genera are numbered according to the list published by De Dalla Torre and Harms in their 
Genera Siphonogamarum (1900-1907) in order to facilitate reference, though genera in the Flora 
are not necessarily arranged in this sequence. 

As in previous volumes, generally accepted abbreviations are used for literature references, 
except in the following cases which appear frequently and are, therefore, considerably condensed: 

C.F.A Conspectus Florae Angolensis 

F.C Flora Capensis 

F.C.B Flore du Congo et du Rwanda-Burundi 

F.S.W.A Prodromus einer Flora von Sudwestafrika 

F.T.A Flora of Tropical Africa 

F.T.E.A Flora of Tropical East Africa 

F.W.T.A Flora of West Tropical Africa 

F.Z Flora Zambesiaca 

Phill., Gen. ed. 2 The Genera of South African Flowering Plants by E. P. 

Phillips, ed. 2 (1951) 

Burtt Davy, FI. Transv Manual of the Flowering Plants and Ferns of the Transvaal 

and Swaziland, Vol. 1 (1926) and Vol. 2 (1932). 

As before, the abbreviation “l.c.” is used for previously cited references even though “op. 
cit.” or “tom. cit.” would in certain cases be more correct. 

In citing specimens the grid reference system has been used. The spelling of the names of 
some localities has been brought into line with the findings of the Committee on Standardisation of 
Place Names. 

In the text, species which show evidence of becoming naturalized are treated in the same way 
as indigenous species. In the Index, synonyms are in italics while exotic species are signified by an 

A change in the existing Flora format is being introduced shortly and in future families or 
sections of large families will be published separately as they are completed. 





P odocarpaceae 









N ajadaceae 



































VOL. 1 

VOL. 2 
VOL. 3 

VOL. 4 

VOL. 5 
VOL. 6 

VOL. 7 
VOL. 8 

VOL. 9 

VOL. 10 













VOL. 11 



VOL. 12 














VOL. 13 








VOL. 14 


VOL. 15 









VOL. 16 

Fabaceae: (1) Mimosoideae 

VOL. 17 




VOL. 18 















VOL. 25 


VOL. 19 


VOL. 26 















VOL. 20 







VOL. 27 






VOL. 28 



VOL. 21 











VOL. 29 

VOL. 22 




VOL. 30 







Turner aceae 


A chariaceae 












VOL. 31 

VOL. 23 












VOL. 24 



VOL. 32 



VOL. 33 





by P. C. V. du Toit and A. A. Obermeyer 

Trees, shrubs or rarely herbs; bark rough or smooth, sometimes flaking; with cortical vascular 
bundles in stem; in Brackenridgea with a yellow pigment beneath bark. Leaves alternate, simple 
(rarely compound in genera outside Africa), discolorous, midrib prominent, nerves penmnerved 
and with reticulate veining, margin entire or serrate; stipules free or united and intrapetiolar, entire 
or laciniate, early caducous or persistent. Flowers bisexual, regular (rarely zygomorphic in genera 
outside Africa), often fragrant, in terminal or axillary panicles, racemes or fascicled cymes, or 
solitary; pedicels articulated at or near base. Sepals usually 5 (3-10), free or nearly so, quincuncial, 
usually persistent and accrescent and often red coloured in fruit. Petals usually 5 (10), contorted in 
bud, fugaceous. Stamens few to many, free; (staminodes sometimes present in genera outside 
Africa); filaments free, persistent; anthers bilocular, basifixed, linear, opening by longitudinal slits 
or apical pores, usually caducous. Carpels (2) 5-15, superior, free, uni-ovulate, or fused, with 
2-many biseriate ovules in each locule; styles as many as placentas, gynobasic or epigynous, 
completely united or free apically, stigmas terminal, usually somewhat enlarged. Fruit: drupes on 
an enlarged torus, berries or septicidal capsules; seeds 1-many, with or without endosperm. 

Genera about 30, species about 250, pantropical, extending furthest south in subtropical and temperate southern Africa. 
Two genera and 13 species occur in our area. 

Leaf margin serrulate with mucronate teeth or more or less entire; lateral veins not continuing 

upwards alongside margin; petals yellow or orange 1 . Ochna 

Leaf margin closely beset with small oblong-ovoid, white glands on the shallow teeth; lateral 

veins continuing upwards alongside margin; petals white or pink 2. Brackenridgea 

5112 1 OCHNA 

OchnaL., Sp. PI. 1: 513 (1753); Gen.Pl. ed. 5: 229(1754); DC., Ann. Mus. Paris 17: 410(1811); 
Prodr. 1: 735 (1824); Harv. in F.C. 1: 448 (1860); Oliv. in F.T.A. 1: 316 (1868); Phill. in Bothalia 
1: 87 (1921); Gilg in Pflanzenfam. ed. 2,21: 53 (1925); Burtt Davy, FI. Transv. 1: 237 (1926); 
Robson in F.Z. 2: 225 (1963); Schreiber in F.S.W.A. 42: 2 (1968). Type species: O. jabotapita L. 
from Sri Lanka. 

Diporidium Bartl. & Wendl., Beitr. Bot. 2: 24 (1825). 

Trees or shrubs or occasionally virgate, soboliferous shrublets; bark rough or smooth, 
sometimes flaking. Leaves semi-persistent, shed in spring or in times of drought, usually shortly 
petiolate, simple, ovate, obovate, oblong to linear, margin entire to serrate, midrib prominent on 
both sides, veins spreading, distinctly reticulate; stipules in S. Afr. spp. intrapetiolar, fused, bifid or 
bidentate. Flowers bisexual, regular, often fragrant, in terminal or axillary panicles or racemes or 
reduced to subumbellate, many- to few-flowered cymes or solitary, on short lateral spurs; bud 
scales or bracts distichous, caducous, leaving transverse scars on twigs; pedicels articulated near 
the base or in lower half, accrescent in S. Afr. spp. Sepals 5 in S. Afr. spp., quincuncial in bud, 
green or yellow, persistent, enlarging and usually turning red or pink in fruit, firm, enveloping bud 



and usually young fruit, but spreading at anthesis and when fruit is ripe. Petals 5 (in S. Afr. spp.), 
yellow, orange or rarely white, usually obovate, attenuate or clawed at the base, fugaceous. 
Stamens many (30-50), free, irregularly placed in a dense whorl on torus; filaments filiform, 
persistent, apex slightly capitate; anthers bilocular, opening by longitudinal slits or apical pores, 
basifixed, caducous. Carpels 5-15, free, attached basally or laterally to the receptacle; styles 
gynobasic, completely united or the upper ends free, recurved; stigmas apical, as many as carpels 
or fused and capitate. Drupelets 1-several, some usually aborting, kidney-shaped and attached 
centrally at the indented inner side or oblong-globose, attached basally to the swollen, often red 
receptacle, usually hard, black and shiny. 

Species about 86, found in Africa south of the Sahara, Yemen, Madagascar, Mascarenes to Asia; tropics. Twelve 
species in Southern Africa, widespread but absent from the Winter-rainfall Region. 

The fruit is distributed by birds who eat the oil-rich drupelets. This could explain the often erratic distribution of 
species. Of interest is the occurrence of 7 taxa in a small area in the N.E. Transvaal on the farm Cyprus, where, because of 
congenial conditions, Ochna arborea vars. arborea and oconnorii, O. confusa, O. holstii, O. natalitia, O. pretoriensis and 
O. serrulata were all well established. 

Galls are found on Ochna natalitia, O. serrulata, O. gamostigmata and O. barbosae. Their shapes are typical for the 
respective species. The following species are soboliferous: O. pulchra, O. confusa, O. natalitia, O. gamostigmata and O. 

Flowers arranged in many-flowered inflorescences or in reduced few-flowered, subumbellate fascicles: 

Carpels and drupelets kidney-shaped, attached centrally to the receptacle on the indented side; bark flaking to 
expose patches of a different colour: 

Carpels 7 or 8; leaves elliptic c. 10 x 4 cm, firm; racemes in simple, many-flowered, pendulous 

bunches 1. O. pulchra 

Carpels 5; leaves narrowly oblong, c. 6 X 1,5 cm (larger in forest trees), thin; racemes compound, many-flowered, 

to short, subumbellate, few-flowered clusters 2. O. arborea 

Carpels and drupelets oblong-globose, attached basally to receptacle; bark not flaking, evenly coloured: 

Anthers opening by longitudinal slits; carpels 5—6: 

Usually arborescent with the leaves placed in a horizontal plane, appearing almost digitate on the ends of the 
branchlets; leaves acute basally, attenuate-acuminate distally; pedicels articulated above the 

base 3. O. holstii 

Usually soboliferous virgate dwarf shrubs with the bark of the woody stem-base not properly developed on 
one side; leaves long-attenuated basally, acute distally; pedicels articulated at the 

base 4. O. confusa 

Anthers opening by apical pores; carpels 6-12 5. O. natalitia 

Flowers solitary (rarely 2 placed close together) on short lateral spurs; carpels 5(6), oblong-globose, attached basally to 

Sepals 1—1,5 cm long in fruit: 

Leaves green, flat and spreading; petioles 1-2 mm long, stout: 

Pedicels erect or spreading, 1-1,5 cm long, articulated at or near base: 

Young twigs with some scattered lenticels; leaves narrowly obovate; margin indistinctly to distinctly 

serrulate; Transvaal 6. O. pretoriensis 

Young twigs conspicuously and densely pale-mottled with lenticels; leaves narrowly elliptic or linear- 
acuminate; margin sharply and closely serrulate: 

Leaves narrowly elliptic; style with 5 free, recurved apical ends bearing small discoid stigmas; pedicels 

articulated 1-2 mm above the base; E. Cape, Natal, E. Transvaal 7. O. serrulata 

Leaves linear-acuminate; style ending in a capitate stigma; pedicels articulated at the base; Natal, E. 

Transvaal 8. O. gamostigmata 

Pedicels recurved, 2-3 cm long, articulated below centre with upper part often forming an angle with lower 

part 9. O. inermis 

Leaves glaucous, drooping and folded; petioles 4 mm long, thin; sprawling shrubs in granite rock fissures; N. 

Transvaal 10. O. glauca 

Sepals 1,5-2 cm long in fruit: 

Virgate, soboliferous shrubs up to 1 m high; young twigs densely lenticellate; sepals loosely arranged around 

maturing fruit; northern S.W. Africa 11. O. cinnabarina 

Trees or large shrubs; young twigs with few lenticels; sepals forming a closed, ovoid covering around maturing 

fruit, spreading later; coastal dunes of Natal 12. O. barbosae 



1. Ochna pulchra Hook., Ic. PI. 6, t.588 
(1843), as pulchrunr, Harv. in F.C. 1: 449 
(1860); Oliv. in F.T.A. 1: 317 (1868); Gilg in 
Bot. Jahrb. 33: 234 (1903); Phill. in Bothalia 1: 
91 (1922); Burtt Davy, FI. Transv. 1: 238 
(1926); Verdoorn in Flow. PI. Afr. 29: 1. 1139 
(1952); Letty, Wild Flow. Transv. 217, 1. 109, 
f.l (1962); Story in Mem. Bot. Surv. S. Afr. 30: 
35, photo. 33 (1958); Robson in F.Z. 2: 228 
(1963); Schreiber in F.S.W.A. 42: 2 (1968); 
Palmer & Pitman, Trees S. Afr. 3: 1503 ( 1973). 
Type: Transvaal, Magaliesberg, Burke & 
Zeyher 191 (K, holo.; PRE, photo.; SAM!). 

O.rehmannii Szyszyl., Polypet. Disc. Rehm. 2: 28 
(1888). Type: Transvaal, Pretoria, hills above Apies River, 
Rehmann 4341 (Z, holo.; K; PRE, photo.). O. ascher- 
soniana Schinz in Verh. Bot. Ver. Prov. Brandenb. 29: 61 
(1888). Type: S.W. Africa, Ovamboland, Otjiheveta, 
Schinz 1139 (Z, holo.; K; PRE, photo.). O. fuscescens 
Heine in Mitt. Bot. Staatssamml. Miinchen 1: 340 (1953). 
Syntypes: S.W. Africa, Waterberg Plateau, Volk 1 1 35(M); 
farm Rotenfels, Rehm s.n.(M). 

Trees up to 10 m tall or shrubs, sobolifer- 
ous, occasionally dwarfed; trunk with rough 
broken bark below, smooth above, grey, flaking 
to expose cream patches. Branches spreading, 
brittle, rough, grey, new growth pale fawn. 
Leaves semi-persistent, turning yellow in wint- 
er, shed in spring, young leaves appearing with 
flowers, oblong to elliptic, c. 10 cm long and c. 
2 cm wide but variable in size, apex obtuse, 
apiculate, base rounded or somewhat attenuate, 
margin pale, entire in lower half, shallowly 
serrulate above, midrib and veins prominent, 
firm and glossy; stipules intrapetiolar, paired, 
subulate, up to 15 mm long, rarely narrowly 
spatulate at the apex, early deciduous. Flowers 
fragrant, in simple, pendulous, clustered, 
many-flowered racemes terminating short an- 
nual branchlets; pedicels c. 1 cm at anthesis, up 
to 1,8 cm in fruit, articulated above the base. 
Sepals broadly ovate, 8 mm long, green to 
yellow, enlarged, reflexed, pink or red and 
white-edged in fruit. Petals soft yellow, 
obovate-attenuate, 12 mm long, fugaceous. 
Stamens numerous (c. 50) with filaments 2 mm 
long; anthers 1 mm long, biporose. Carpels 
6-8, attached centrally on inside of torus; styles 
fused below, free and recurved above; stigmas 
apical, tumid. Drupelets kidney-shaped, 
rounded, 12 mm long, black or mottled with 
cream. Fig. 1: 1; 2: 1. 

Recorded from the Transvaal, South West Africa, 
Angola, Rhodesia, Zambia and western Mozambique; form- 
ing colonies on rocky sandstone slopes or in sandy areas. 
Flowering in September-October. 

New growth from stolons in narrow rock fissures may 
remain hemicryptophytic and has on occasion been mis- 
taken for the poisonous Dichapetalum cymosum. 

S.W. A. — 1715 (Ondangua): Oshikango near Angola 
border, Rodin 2615; 2662. 1718 (Kuring-Kuru): 12 km E. 
of Kuring-Kuru, Giess 9504. 1722 (Chirundi): Western 
Caprivi Strip, Bwabwata Rest Camp, Watt 18. 1918 (Groot- 
fontein): N.E. of Grootfontein, Schoenfelder 220. 1919 
(Kanovlei): Okavango Territory, Kanovlei, Barnard 4. 
1920 (Tsumkwe): Garu, 88 km S. of Runtu, Maguire 1576. 
2012 (Waterberg): Waterberg Plateau, Boss sub TRV 

Transvaal. — 2230 (Messina): Makonde, 24 km E. of 
Sibasa, Pienaar 47. 2231 (Pafuri): Punda Milia, Van der 
Schijff 947; 996. 2427 (Thabazimbi): Rankin’s Pass, Van 
Wyk 42 ; near Rooiberg, Codd 6154. 2428 (Nylstroom): 13 
km. N. of Nylstroom on road to Naboomspruit, Vahrmeijer 
134; Maguire 1354; Mosdene, Galpin M52. 2429 
(Zebediela): Potgietersrus, Leendertz 1258. 2526 (Marico): 
Zeerust, Thode A 1367. 2527 (Rustenburg): Rustenburg, 
Galpin 11633; 11635 ; Magaliesberg, Pegler 1032; Har- 
tebeestpoortdam, Gerstner 6416. 2528 (Pretoria): 

Meintjieskop, Mogg 15310; Burtt Davy 5035. 2529 (Wit- 
bank): Loskop Dam area, Prosser 1885; Mogg 30579. 2627 
(Potchefstroom): Jack Scott Reserve, near Krugersdorp, 
Wells 2338. 

Common names: Lekkerbreek; Monyelenyele (Tsw); 
Monamane, Mopha (NS); Musuma (V). 

2. Ochna arborea Burch, ex DC., Prodr. 
1: 736 (1824). Type: E. Cape, Port Alfred, 
Kowie River, Burchell, Cat. 4012 (G, holo.; K; 
PRE, photo.). 

Understorey forest trees up to 12 m tall, or 
smaller shrubs; trunk with light red wood, 
covered with smooth grey bark, flaking irregu- 
larly exposing brown patches; crown small, 
with ascending grey and brown mottled 
branches which are irregularly ridged with grey 
raised lenticels. Leaves semi-persistent or de- 
ciduous, shed in spring or during a period of 
drought; lamina deep green, variable, broadly to 
narrowly elliptic, 5-10 cm long, 2-3 cm broad, 
apex rounded to acute, base truncate, margin 
nearly smooth or with some shallow teeth to 
sharply serrate, venation distinctly reticulate, 
leathery; stipules intrapetiolar, fused. Flowers 
scented, arranged in compound, complex, 
many-flowered usually erect panicles on well 
developed, mostly leafless branchlets to a re- 
duced subcorymbose fascicle with 4—8 flowers 
on a very short spur. Sepals 5, ovate-convex, c. 
7 mm long, green; enlarged, dark red and 
reflexed at anthesis and in fruit. Petals 5, 
obovate-cuneate, 12 mm long, yellow. Stamens 
c. 30, with filaments about as long as biporose 
anthers. Carpels 5, reniform; styles completely 



Fig. 1. — 1, Ochna pulchra, flowering branch, x l /y t la, old flower showing persistent calyx, torus and 1 drupelet attached 
ventrally, X 1 (ex garden, Bol. Res. Inst.). 2, O. pretoriensis, flowering branch, x 1; 2a, old flower showing 
persistent calyx, torus and 3 drupelets attached basally, x 1 (ex garden, Bot. Res. Inst.)\ 2b, anther with locules 
opening through apical pores, X 10. 3, O. confusa, anther with longitudinal dehiscence, x 10. 4, O. natalitia, gall 
(Moll 3612) x 1. 5, O. serrulata, gall (McClean & Ogilvie sub PRE 39308), x 1. 



united or free apically; stigmas capitate. 
Drupelets reniform, 10-15 mm long, black. 

Transvaal, Swaziland, Natal to eastern and south- 
eastern Cape as far as George; also in Mozambique and 
Rhodesia. Flowering in spring. 

Two varieties are recognized. 

Leaves broadly elliptic, c. 4 X 1,5 cm; marginal teeth 
often obscure; inflorescence few-flowered on 
short spurs; trees to shrublets found outside or 

inside forests (a) var. arborea 

Leaves narrowly elliptic — acuminate, c. 8 x 2,5 cm; 
margin distinctly serrate; inflorescence many- 
flowered, paniculate; forest 

tree . . .(b) var. oconnorii 

(a) var. arborea. 

Ochna arborea Burch, ex DC., Prodr. 1: 736 (1824); J. 
Smith in Bot. Mag. t.45 19 (1850); Harv. in F.C. 1: 449 
(1860); Sim, For. FI. Cape Col. 163, t.29, f.l (1907); Phill. 
in Bothalia 1: 92 (1922); Burtt Davy, FI. Transv. 1: 238 
(1926); Robson in F.Z. 2: 230 (1963); Palmer & Pitman, 
Trees S. Afr. 3: 1504 (1973). Type: E. Cape, Port Alfred 
Kowie River, Burchell, Cat. 4012 (G; holo.; K; PRE, 

Diporidium arborewn (Burch, ex DC.) Wendl. in Bartl. 
& Wendl., Beitr. Bot. 2: 26 (1825). Type as above. 

The variety arborea is distinguished by its 
shorter and relatively broader elliptic leaves, 
which usually have subentire margins. The 
inflorescence is few-flowered. 

It appears to be the form adopted in less favourable 
conditions. The distribution is mainly southern, in forest 
scrub, dune forest and bushveld. Fig. 2: 2a. 

Transvaal. — 2229 (Waterpoort): Wylliespoort, 
Hafstrom & Acocks 1895 . 2430 (Pilgrims Rest): Cyprus, 
Renny 137\ The Downs, Du Toit 178. 

Swaziland. — 2631 (Mbabane): 25 km S. of Stegi, 
Compton 29282. 

Natal. — 2632 (Bela Vista): Ndumu Game Reserve, 
Pooley 870. 2731 (Louwsburg): Sokosoko forest, Nongo- 
ma, Gerstner 4910. 2732 (Ubombo): 21 km from In- 
gwavuma to Ndumu, Moll & Strey 3710\ Mkuzi Game 
Reserve, Ward 3604. 2830 (Dundee): Qudeni Forest, Ed- 
wards 2655. 2831 (Nkandla): Eshowe, Meebold 12952; 
Thode A 1228. 2832 (Mtubatuba): False Bay, Ward 1634. 

2930 (Pietermaritzburg): Karkloof, Lions River, Ehlatini 
Forest, Moll 2862 ; Kloof near Pinetown, Galpin 12080. 

2931 (Stanger): Durban, Franks sub TRV 34099. 3030 
(Port Shepstone): Gibraltar, Strey 10041 . 

Cape. — 3129 (Port St. Johns): Mpingo Forest, Mtamba- 
la, Fegen sub PRF 3089. 3227 (Stutterheim): Kabaku Hills, 
Acocks 8967; Nahoon River, East London, Galpin 3291 . 
3228 (Butterworth): Kentani, Pegler 1216; near Komga, 
Flanagan 359. 3322 (Oudtshoom): Ebb & Flow Nature 
Reserve, Taylor 7984. 3325 (Port Elizabeth): Van Stadens 
Pass, De Winter 7618. 3326 (Grahamstown): Bushmans 
River mouth, Galpin 10667; Port Alfred, Salisbury 68. 
3423 (Knysna): Harkerville Forest, Keet 52 7; Storms River, 
Keet 525. 3424 (Humansdorp): Witelsbos, Fourcade 845. 

Common names: Cape plane, redwood; rooihout; um- 
Tensema (X); umBomvane, umTelele (Z). 

(b) var. oconnorii (Phill.) Du Toit in 
Bothalia 11: 518 (1975). 

O. oconnorii Phill. in Bothalia 1: 92 (1922); Robson in 
F.Z. 2: 230 (1963). Lectotype: Transvaal, Woodbush forest, 
O’Connor sub PRE 1257 (PRE, lecto.; PRF 2198). 

This variety is a more luxuriant forest form 
with longer, larger leaves and erect, compound, 
elongate inflorescences. There are inter- 
mediates. Fig. 2: 2b. 

Recorded from the mist belt forests of Transvaal, Natal 
and eastern Cape; also in Rhodesia and Mozambique. 

Transvaal. — 2230 (Messina): Entabeni Forest Reserve, 
Obermeyer sub TRV 30346. 2330 (Tzaneen): Westfalia 
Estate, Scheepers 728; Woodbush, Botha 1; Grenfell 3. 
2430 (Pilgrims Rest): The Downs, Du Toit 178; Pilgrims 
Rest, Burtt Davy 1409. 2531 (Komatipoort): Barberton, 
Oranje sub PRE 1264; Thorncroft 2029. 

Natal. — 2632 (Bela Vista): Ndumu Game Reserve, 
Pooley 135c. 2831 (Nkandla): Eshowe, Moberly 9. 2930 
(Pietermaritzburg): York, Benvie, Fisher 1038. 

Cape. — 2326 (Grahamstown): The Haven, Gordon-Gray 

Common names: Cape Plane, Rooihout, Rooi yster- 
hout, Morelle; Murambo, murambothavha (V). 

3. Ochna holstii Engl, in Abh. Preuss. 
Akad. Wiss.: 69 (1894); Pflanzenw. Ost-Afr. C: 
273 (1895); Gilg in Bot. Jahrb. 33: 234, 241 
(1903); Phill. in Bothalia 1: 93 (1922); Burtt 
Davy, FI. Transv. 1: 238 ( 1926); Robson in F.Z. 
2: 240 (1963); Palmer & Pitman, Trees S. Afr. 
3: 1507 (1973). Type: Tanzania, Usambara, 
Holst 2601 (Bf; K; PRE, photo.). 

Understorey trees up to about 8 m tall or 
shrubs with red wood and rough ridged, grey or 
brown bark which does not flake; branches 
spreading; young branchlets with scattered 
raised lenticels, glabrous or puberulous. Leaves 
tardily deciduous, immature at time of flower- 
ing, radiating in a horizontal plane from the tips 
of young shoots in a somewhat digitate pattern; 
blade narrowly elliptic, 5-10 cm long, 1, 5-2,5 
cm broad, apex attenuate-acuminate, base 
cuneate, regularly and closely serrulate, stiffly 
membranous, lateral veins curving upwards 
from midrib; stipules intrapetiolar, fused. Flow- 
ers usually in (2) 6-10-flowered racemes, on 
short lateral spurs below new leafy shoots 
(tropical forms with many-flowered elongate 
racemes); pedicels thin, up to 3 cm in fruit, 
articulated above base, glabrous or puberulous. 



Kig. 2. — I, Leaves of South African species of Ochna x 1: I, O. pulchra ( Wells 2031). 2a, O. arborea var. arborea 
(Archibald 6125); 2b, var. oconnorii (Wager sub TRV 22971). 3, O. holstii (Gersrner 5998). 4, O. confusa (Remix 
138). 5, O. natalitia (Galpin 13569). 6, O. pretoriensis (Smith 1239). 7, O. serrulata (Moll 2510). 8. 6. 
gamostigmata (Moll 2861). 9, O. inermis (Codd 4824). 10, O. glauca ( Verdoorn 2292). 11, O. cinnabarina 
(Killick <£ Leistner 3202). 12, O. barbosae (Balsinhas 1372). 



Sepals narrowly elliptic, c. 9 mm long, obtuse, 
ivory at anthesis, up to 1,5 cm long, red, pink, 
purple or green and spreading in fruit, some- 
what crinkled when dried. Petals obovate, 
about 1 cm long, clawed, pale to bright yellow, 
caducous. Stamens c. 40, with filaments 3 mm 
long; anthers 1,5 mm long, dehiscing by lon- 
gitudinal slits. Carpels 5(-6), erect; style with a 
swollen, lobed stigma. Drupelets oblong- 
globose, c. 8 mm long, attached basally to 
swollen torus. Fig. 2: 3. 

An understorey forest tree recorded from the Trans- 
vaal, Natal and eastern Cape; widespread and variable in its 
range to Rhodesia and tropical eastern Africa; inhabiting 
shady ravines in mistbelt forests, gallery forests along rivers 
or as a forest relic in open areas; flowering October - 

Transvaal. — 2229 (Waterpoort): Soutpansberg above 
Louis Trichardt, Hutchinson 2025; Story 5935. 2230 (Mes- 
sina): Soutpansberg, Phiphidi Waterfall near Sibasa, Ober- 
meyer sub TRV 30037; Lake Funduzi, Story 4850. 2329 
(Pietersburg): Lejuma Plateau, Strey 7945; Nicholson 669. 
2330 (Tzaneen): Woodbush, Botha sub PKF 2935. 2428 
(Nylstroom): near Palala, Smuts & Gillett 3357. 2430 
(Pilgrims Rest): Cyprus, Renny 186; Mariepskop, Van der 
Schijff 4754; 5918; 6237. 2527 (Rustenburg): kloof in 
western Magaliesberg range. Rose Innes 203; near Beacon 
6078, Taylor sub PRE 28763. 2530 (Lydenburg): Kaapse 
Hoop, Berlin Forest Reserve, Hofmeyr sub PRE 28762. 
2531 (Komatipoort): Barberton, 5 km S.W. of Agnes Mine, 
Story 5993; 5455. 

Natal. — 2731 (Louwsburg): Ngome Forest Reserve, 
Gerstner 4496; 5152; Tustin sub PRF 3115. 2830 (Dun- 
dee): Qudeni Forest Reserve, Edwards 2667. 2930 (Pieter- 
maritzburg): Noodsberg, Ward 938. 3029 (Kokstad): Ingeli 
Forest Reserve, Tustin sub PRE 2907. 

Cape. — 3127 (Lady Frere): Engcobo Mt., Flanagan 

Common names: Rooi Ysterhout; Real Red Pear; 
isiBanku, Pambane (Z); Tshipfure (V); ngqelene (X). 

4. Ochna confusa Burtt Davy & Green- 
way in Burtt Davy, FI. Transv. 1: 238 (in 
clavis), 239 (1926); in Kew Bull. 1926: 239 
(1926); Robson in F.Z. 2: 250 (1963). Syn- 
types: Transvaal, Barberton, Rogers 18264 
(PRE); Pilgrims Rest, Rogers 23068 (PRE). 

O. leptoclada sensu Phill. in Bothalia 1: 94 (1922) pro 
parte quoad specim. Transvaal. 

Saxicolous, soboliferous shrubs or dwarf 
shrubs 0,5— 1(— 2) m tall, the virgate branches 
arising from a woody base; this basal sprawling 
stem lacking bark on one side (in Transvaal 
specimens seen). Branches and branchlets pale 
dirty grey, lenticellate, rough. Leaves decidu- 
ous, sessile, narrowly elliptic, 6-9 cm long, 
1,5-2, 5 cm broad, apex attenuate, acute or 

obtuse, base long-attenuate, margin serrulate, 
firm, with close lateral veins curving upwards; 
stipules intrapetiolar, elongate-deltoid. Flowers 
in sessile (1-) 2-6-flowered pseudumbels on 
short lateral spurs below new leaf growth; 
pedicels about 12 mm long in fruit, articulated 
at the base. Sepals c. 5 mm long, ovate at 
anthesis; 1 cm long, deep red, flat and spreading 
in fruit. Petals obovoid, clawed, c. 1 cm long, 7 
mm broad, bright yellow. Stamens c. 25-30, 
with filaments 5 mm long; anthers 2 mm long, 
opening by longitudinal slits. Carpels 5, at- 
tached basally to receptacle; styles fused, terete; 
stigma capitate. Drupelets oblong-globose, c. 1 
cm long, black. Fig. 1: 3; 2: 4. 

Recorded from eastern Transvaal; also in Mozam- 
bique, Malawi, Rhodesia, southern Tanzania, Zambia. In- 
habits rock crevices on hillsides in more open vegetation; 
flowering in spring. 

Transvaal. — 2430 (Pilgrims Rest): farm Cyprus, Renny 
138; Sikorora near Macoutsie River, Van Darn sub TRV 
22935; Ohrigstad Nature Reserve, Jacobsen 1242; Graskop 
Falls, Galpin s.n. 2530 (Lydenburg): Waterval Boven, 
Britten 4741; 4770. 2531 (Komatipoort): Barberton, Nel 
52; Thode A 1570. 

O. confusa shows a relationship with Brackenridgea in 
that its anthers open by longitudinal slits, and in its leaves, 
which have lateral veins curving upwards and marginal 
teeth tipped with glands. 

5. Ochna natalitia (Meisn.) Walp., Re- 
pert. 2: 826 (1843); Phill. in Bothalia 1: 93 
(1922); Burtt Davy, FI. Transv. 1: 239 (1926); 
Robson in F.Z. 2: 233 (1963); Palmer & Pit- 
man, Trees S. Afr. 3: 1508 (1973). Type: Natal, 
Port Natal (Durban), Krauss 454 (K, holo.; 
PRE, photo.). 

Diporidium natalitium Meisn. in Hook., J. Bot. Lond. 2: 
58 (1843). Type as above. 

O. atropurpurea var. natalitia (Meisn.) Harv. in F.C. 1: 
448 (1860). O. chilversii Phill. in Bothalia 1: 90 (1922). 
Syntypes: E. Cape, Ingeli Forest Reserve, Chilvers sub PRF 
1945 (PRE; PRF; K); at mouth of Umkwani River, Tyson 
2619 (PRE). 

Understorey forest trees up to 6 m tall with 
red wood and a dark rough bark not peeling off; 
shrubby in open vegetation or reduced to 
soboliferous shrublets when exposed to fire and 
frost, 0,5-2 m tall. Branches and branchlets 
dark brown or grey, lenticellate; young twigs 
pale fawn coloured, the epidermis peeling off in 
small thin membranous white flakes. Galls 
often present, globose with rosulate, ovate, 
obtuse bracts. Leaves tardily deciduous, nar- 
rowly oblong, variable in size, 5-12 (-14) cm 



long, 1,5-2, 5 cm broad, apex and base gener- 
ally obtuse, margin serrulate, rarely subentire, 
midrib distinct, lateral veins at right angles to 
midrib, leathery, shortly petiolate; stipules in- 
trapetiolar, elongate-deltoid. Flowers fragrant, 
in compound, many- to few-flowered racemes 
or subumbellate, terminating short lateral spurs; 
pedicels up to 2,5 cm long, articulated in lower 
quarter. Sepals elliptic-convex, about 9 mm 
long, green, slightly larger in fruit, turning wine 
red. Petals broadly obovate, c. 15 mm long, 
clawed, yellow. Stamens c. 30, with filaments 3 
mm long; anthers biporose, 2 mm long. Carpels 
6-12, usually about 8, erect; styles fused, terete, 
the apical ends free, recurved, forming swollen 
irregular glandular stigmas. Drupelets oblong- 
globose, 10-15 mm long, attached basally, 
black, shiny. Fig. 1: 4; 2: 5. 

Fairly common along the eastern Cape coastal belt 
from Knysna to Natal and northern Mozambique; further 
inland in Natal and Swaziland to eastern and northern 
Transvaal, in forests or coastal scrub, in sandy soil or 
amongst rocks; flowering in spring. 

Transvaal. — 2230 (Messina): Makonde Mission Sta- 
tion, Westphal sub TRV 29126. 2231 (Pafuri): Kruger 
National Park, Punda Milia, Rowland Jones 27. 2327 
(Ellisras): Soutpan, Galpin 15150. 2329 (Pietersburg): 
Louis Trichardt, Breyer sub TRV 23390; 24385. 2430 
(Pilgrims Rest): Mariepskop, Killick & Strey 2440. 2431 
(Acomhoek): near Skukuza, Cholmondely s.n. 2528 (Pre- 
toria): Kameeldrift, 64 km N.E. of Pretoria, Repton 3875. 

2529 (Witbank): Loskopdam, Tweefontein, Theron 1512. 

2530 (Lydenburg): Nelspruit, Breyer sub TRV 17701; 
Buitendag 323; 226. 2531 (Komatipoort): Pretoriuskop, 
Van der Schijff 6. 

Swaziland. — 2531 (Komatipoort): Komassan River, 
Havelock Concession, Saltmarshe sub Galpin 983. 2631 
(Mbabane): Hill N.E. of Mbabane, Compton 26184; Nton- 
dozi, Compton 29211. 

Natal. — 2632 (Bela Vista): Ndumu Game Reserve, 
Pooley 135b. 2731 (Louwsburg): Nongoma, Gerstner 
3642; 5105. 2732 (Ubombo): Sihangwane Store, Moll 
4400; Mkuzi Game Reserve, Ward 3611. 2831 (Nkandla): 
Eshowe, Gerstner 2818; Nkandla Forest, Edwards 2663. 
2832 (Mtubatuba): 25 km N. of Mtubatuba, Codd 2018. 
2930 (Pietermaritzburg): “The Start”, near Howick, Moll 
3370. 2931 (Stanger): Durban, Marloth 4318; Medley 
Wood 66; 12702. 3030 (Port Shepstone): near Southbroom, 
Codd 9706. 

Cape. — 3129 (Port St. Johns): Ntsubane, Strey 9006; 
Notinsila Forest, Fegen sub PRF 1816; Manteku, Dakane 
Location, Strey 10197. 3325 (Uitenhage): Groendalkloof, 
below dam. Mauve 4934. 

Common names: Coast Boxwood, Coast Redwood; 
umBomvu, umBomvane, umMilamatsheni (Z). 

Ochna natalitia and O. arborea are often difficult to 
distinguish in the herbarium when the specimens consist of 
sterile branchlets only and offer no information on bark, etc. 
Galls are found only in O. natalitia. Young twigs of this 
species are fawn coloured. Mature leaves from forest trees 

of both species are usually indistinguishable for both 
become narrowly oblong with coarse marginal teeth, pos- 
sess spreading lateral veins and a dense reticulation. More 
often the lamina of O. natalitia is narrowly oblong whereas 
that of O. arborea is ovate and lighter coloured below. 

In the sandy area of Tongaland, in northern Natal, 
north of Manzengwenya in a strip a few km inland from the 
sea, a form of Ochna natalitia occurs, which is from 7,5 to 
15 cm tall. These plants can cover an extensive area and it 
was found that the individual plants were interconnected by 
underground stems. At first thought to be a new species, 
investigation showed that morphologically they differ from 
the shrub or tree-like form only in the much smaller growth 
form and in having underground stems, a feature in com- 
mon with a few other Ochna species, e.g. O. pulchra, O. 
cinnabarina and O. confusa. The anatomy has proved these 
underground interconnecting parts to be stems, in no way 
differing from the aerial stems. 

6. Ochna pretoriensis Phill. in Flow. PI. 
S. Afr. 2: t.70 (1922); in Bothalia 1: 95 (1922); 
Burtt Davy, FI. Transv. 1: 239 (1926). Syn- 
types: Transvaal, Pretoria, Fouche sub PRE 
1491 (PRE); Phillips sub PRE 1422 (PRE). 

Shrubs or very rarely trees, much 
branched. Stems and branches with a rough 
brown bark, flaking in thin, small, longitudinal 
strips. Leaves tardily deciduous, obovate to 
narrowly obovate or oblong, usually about 3,5 
cm long and 1-1,5 cm broad, apex acute to 
obtuse, base cuneate, margin shallowly serru- 
late with mucronate teeth, midrib occasionally 
arched with the convex half of lamina some- 
what larger than other side, dark green, shiny; 
petiole 1-2 mm; intrapetiolar stipules 
elongate-deltoid, fused at the base. Flowers 
fragrant, solitary or rarely some 2-3-nate, on 
short spurs, appearing before the leaves; 
pedicels c. 8 mm at anthesis, up to 15 mm and 
erect or spreading in fruit; articulated at or near 
the base. Sepals elliptic, c. 5 mm long, obtuse, 
about 14 mm long and wine red in fruit, loosely 
enveloping drupelets. Petals broadly obovate, 
clawed, c. 14 mm long, rounded at the apex, 
yellow. Stamens c. 40, with the filaments 3 mm 
long; anthers biporose, 2 mm. Carpels 5, at- 
tached basally to torus; styles fused with free 
apical recurved tips; stigmas apical, discoid. 
Drupelets oblong-globose, c. 9 mm long, black. 
Fig. 1: 2; 2: 6. 

Endemic, gregarious and widespread in the Transvaal 
with one record from near Mafeking; saxicolous; common 
on northern hill-slopes around Pretoria; flowering in spring. 

Transvaal. — 2229 (Waterpoort): farm Soutpan 193, 
lower northern slopes of Soutpansberg, Obermeyer, 
Schweickerdt & Verdoorn 312. 2429 (Zebediela): granite 
ridge near Utrecht hills, Acocks 8804. 2527 (Rustenburg): 
Buffelspoort, Turner 46. 2528 (Pretoria): Hammanskraal, 



Hutchinson & Mogg 2875; Magaliesberg above Wonder- 
boom Nature Reserve, Dyer 3131; Meintjies Kop, Pole 
Evans 353. 2529 (Witbank): Olifants River Gorge, Mogg 
22422; Doomkloof, Du Plessis 168. 2730 (Vryheid): Piet 
Relief, Leipoldt sub PRE 30203. 

Cape. — 2525 (Mafeking): Blikplaas, Mafeking district, 
Brueckner 472. 

Related to O. inermis but can be distinguished by the 
shorter usually erect pedicels articulated just above the base, 
and the somewhat smaller leaf. 

7. Ochna serrulata (Hochst.) Walp., Re- 
pert. 5: 400 (1846); Du Toit in Bothalia 11: 517 
(1975). Type: Natal, Natal Bay (Durban), 
Krauss 473 (Bf; K; PRE, photo.). 

Diporidium serrulatum Hochst. in Flora 27, 1: 304 
(1844). Type as above. 

O. atropurpurea sensu Harv. in F.C. 1 : 448 ( 1860); sensu 
Sim, For. FI. Cape Col. 163, t. 29, f.2 (1907); sensu Phill. in 
Bothalia 1: 94 (1922); sensu Stapf in Bot. Mag. t.9042 
(1925); sensu Burtt Davy, FI. Transv. 1: 239 (1926); sensu 
Van der Watt in Flow. PI. Afr. 35: 1. 1392 (1962); sensu 
Palmer & Pitman, Trees S. Afr. 3: 1509 (1973), non DC. O. 
multiflora Williams in J.R. Hort. Soc. 5: 160 (1897), nom.; 
Gard. 22: 574, t.369 (1882); Gard. Chron. 40: 212, icon, 
suppl. (1906); non DC. 

Decorative evergreen shrubs, 60-200 cm 
tall, or rarely a small tree. Stems with red wood; 
branches brown, densely verrucose with pale 
grey, raised rounded lenticels. Galls often pre- 
sent, forming a globose body consisting of 
numerous subulate bracts. Leaves partly shed at 
time of flowering, narrowly elliptic, 2,5-5 cm 
long, 1-1,5 cm broad, apex and base obtuse to 
acute, margin sharply serrulate with the teeth 
and apex mucronate, thinly coriaceous, shiny, 
dark green, midrib distinct, reticulate-veined; 
petiole 1 mm; intrapetiolar stipules linear, 
bidentate, margin toothed. Flowers usually sol- 
itary or rarely binate, on short lateral spurs; 
pedicels 1 cm long at anthesis, 1,5 cm in fruit, 
articulated 1-2 mm above base. Sepals ovate, 8 
mm long at anthesis, green, up to 15 mm long 
and deep wine red in fruit, spreading or 
reflexed. Petals broadly obovate, cuneate, 1 cm 
long. Stamens c. 20, with filaments c. 3 mm 
long; biporose anthers about equal in length. 
Carpels 5, attached basally; style terete with 
short, recurved tips; stigmas apical, discoid. 
Drupelets oblong-ovoid, c. 1 cm long, black. 
Fig. 2: 7. 

Recorded from the eastern Transvaal to Natal and 
eastern Cape as far west as George; common along forest 
edges or beside paths, in scrub forest or fynbos. Flowering 
in spring, but with some flowers throughout the year. 

Schlechter collected specimens at Genadendal in 1896 
but there are no further records from this area; it is possible 
that the plants were introduced at the Mission Station. 

Transvaal. — 2430 (Pilgrims Rest): road from the 
Downs to Madeira, Crundall sub PRE 32796. 2730 (Vry- 
heid): Mooihoek, near Piet Retief, Devenish 1264. 

Swaziland. — 2631 (Mbabane): Hlatikulu, Galpin 9622. 

Natal. — 2731 (Louwsburg): Ingwavuma, Ward 2374. 
2829 (Harrismith): Cathedral Peak Area, Inhlwasine River 
Valley, Killick 1816; Pieters, Strey 9545. 2831 (Nkandla): 
Melmoth, Mogg 6142; Eshowe, Gerstner 2031. 2929 
(Underberg): Tabamhlope Research Station near Estcourt, 
West 817; 531. 2930 (Pietermaritzburg): Albert Falls, 
Meebold 12948; Thomville, Moll 3418. 2931 (Stanger): 
Groutville, Moll 2510; Durban, Natal Bay, Gueinzius s.n. 
3029 (Kokstad): 10 km S. of Umzimkulu on Harding Road, 
Story 655. 3030 (Port Shepstone): Dumisa, Rudatis 667; 
Horseshoe farm, Strey 5872. 

Cape. — 3128 (Umtata): Umtata commonage, Miller 
B/952. 3129 (Port St. Johns): Second Beach, Bruce 448; 
Prospect Siding, Flanagan 322. 3226 (Fort Beaufort): 
Katberg forest. Staples sub PRF 2937. 3227 (Stutterheim): 
Thomas River, Compton 19288; Keiskamma Hoek, Story 
2795; Stutterheim, Rogers 12726. 3228 (Butterworth): 
Qora Mouth, Meeuse 9695; Komga, Flanagan 322; Ken- 
tani, Pegler 194. 3322 (Oudtshoom): Wilderness, Compton 
15565. 3323 (Willowmore): The Crags, Compton 23593. 
3324 (Steytlerville): Melkhoutboom road. Story 2608; hills 
N. of Patensie, Taylor 1283. 3325 (Port Elizabeth): Enon, 
Thode A1099; Suurberg, Swartehoogte, Zeyher sub SAM 
38391. 3326 (Grahamstown): Howiesonspoort, MacOwan 
1766; Grahamstown, Galpin 48. 3419 (Caledon): Gena- 
dendal, Schlechter 7749. 3423 (Knysna): Knysna, Keet 
522; 590. 

The species is occasionally planted as an ornamental 
shrub in gardens in South Africa and in England. At 
Kirstenbosch it is used as a hedge. 

Common names: Rooihout; iLitye (X); umBomvane 

According to the Gardener’s Chronicle (September 
1906), it was first introduced to England in 1820 under the 
name of Ochna multiflora, a catalogue name (not of DC). It 
was lost to cultivation but re-introduced later. In Flowering 
Plants of Africa, 1. 1392 (1962), Ochna serrulata is illus- 
trated (as O. atropurpurea) from shrubs flowering and 
fruiting freely in the garden of the Royal Horticultural 
Society at Wisley, England, in the early 1960s. 

Suspected hybrids: Some specimens collected in Natal 
and the S.E. Transvaal appear to be intermediate between 
Ochna natalitia and O. serrulata. Like O. natalitia they 
bear (few-flowered) umbels, have larger leaves and are 
sometimes arborescent. On the other hand they possess the 
dense mottled covering of lenticels so typical of O. ser- 
rulata and, in fact, anatomically they agree with the latter 
and not with O. natalitia. 

8. Ochna gamostigmata Du Toit in 
Bothalia 11: 517 (1975). Type: Transvaal, Bar- 
berton, Upper Moodies, Galpin 963 (PRE, 
lecto; NH; SAM). 

O. atropurpurea DC. var. angustifolia Phill. in Bothalia 
1: 95 (1921). Syntypes: Transvaal, Barberton, Galpin 963 
(PRE; NH; SAM); Ermelo, Nooitgedacht, Pott 5096 (PRE); 
between Pilgrims Rest and Sabie, Rogers 23410; Swazi- 
land: Havelock Concession, Saltmarshe in Herb. Galpin 
sub PRE 1270 (PRE). 



Small shrubs or rarely trees up to 8 m tall, 
forming suckers. Stems and branches brown, 
densely covered with grey, raised, rounded 
lenticels. Galls sometimes present, forming 
globose bodies consisting of numerous linear- 
subulate bracts spirally arranged. Leaves shed 
at time of flowering, narrowly ovate- to linear- 
acuminate, 2-3 cm long and c. 8 mm broad, 
apex acuminate, rarely acute to obtuse, base 
rounded or acute, margin sharply serrulate with 
the teeth and apex nrucronate, thinly coriace- 
ous, shiny, midrib and lateral veins distinct; 
stipules intrapetiolar, fused, linear, bidentate. 
Flowers sweetly scented, solitary, on short 
spurs; pedicels 1 cm at anthesis, 2 cm long and 
recurved in fruit, articulated at the base. Sepals 
ovate, 7 mm, green, somewhat enlarged in fruit 
and turning brownish red. Petals obovate, 7 
mm, yellow. Stamens 15-25, with filaments 1 
mm long; anthers biporose, 3 mm long. Carpels 
5, attached basally to receptacle; style terete; 
stigma entire, capitate, with short swollen 
lobes. Drupelets oblong-globose, 8 mm Jong, 
black. Fig. 2: 8. 

Recorded from eastern Transvaal, Swaziland and 
Natal, usually at fairly high altitudes, in forested areas. 
Flowering in spring but with some flowers throughout the 

Transvaal. — 2430 (Pilgrims Rest): 17 km E. of Gras- 
kop, Codd & De Winter 3119. 2531 (Komatipoort): Shiya- 
lo-ngubu valley, Pole Evans 4680. 

Swaziland. — 2531 (Komatipoort): Mtutusi River, 

Havelock, Compton 29126. 

Natal. — 2730 (Vryheid): Inkamana Farm, Gerstner 
3516. 2830 (Dundee): Impati Hill, Shirley 13. 2831 
(Nkandla): Eshowe, Thode A1227. 2929 (Estcourt): 

Griffin’s Hill, Acocks 10471. 2930 (Pietermaritzburg): 
Karkloof forest. Farm Ehlatini, Moll 2861; 3375; 3476; 24 
km N.W. of New Hanover, Codd 1465. 

O. gamostigmata differs from O. serrulata in its 
smaller, usually shrubby habit, smaller and narrower, usu- 
ally acuminate leaves, smaller flowers and capitate stigma. 
It grows at fairly high altitudes from Karkloof to Swaziland 
and the eastern Transvaal (Letaba district). 

9. Ochna inermis (Forsk.) Schweinf. 
apud Penzig in Atti Congr. Bot. Intern. Genova 
1892: 335 (1893); Gilg in Bot. Jahrb. 33: 247 
(1904); Schweinf., Arab. Pfl. Aegypt. Alger. & 
Yemen 148 (1912); Robson in F.Z. 2: 237 
(1963); Palmer & Pitman, Trees S. Afr. 3: 1510 
(1973). Type: Yemen, Forskal 760 (C, holo.; 
PRE, photo.). 

Euonymus inermis Forsk., FI. Aegypt. -Arab. 204 
(1775). Type as above. 

O. rogersii Hutch., Botanist in S. Afr. 309, 317 (1946), 
nom. nud. 

Spreading shrubs 1-2 m tall, rarely ar- 
borescent. Stems smooth, dark grey, with pale 
reddish wood; side branches lenticellate, 
numerous, short, bearing spurs which produce 
leaves and flowers. Leaves deciduous, being 
shed just before flowers appear (soon wilting 
after being picked), elliptic, 2-4 cm long and 
1-1,5 cm broad, apex obtuse to acute, base 
acute or rounded, margin shallowly serrulate, 
the teeth mucronulate, shiny, distinctly reticu- 
late; petiole 2 mm; stipules intrapetiolar, fused, 
bifid with apices filiform. Flowers sweet 
scented, solitary or rarely 2 close together, on 
short spurs; pedicels filiform, 1-1,5 cm long at 
anthesis, firm, wiry and up to 2 cm, red and 
recurved in fruit, articulated in lower quarter. 
Sepals ovate, convex, c. 5 mm long, green at 
anthesis, becoming c. 16 mm long and red in 
fruit, at first enclosing young drupelets, spread- 
ing when ripe. Petals yellow, orbicular, 
clawed, about 8 mm. Stamens c. 50, with 
filaments 3 mm long; anthers biporose, 1,5 mm 
long. Carpels 5, attached basally to receptacle; 
styles fused, slender, apices free, recurved, 
filiform with small discoid stigmas. Drupelets 
oblong-globose, 1 cm long, black. Fig. 2: 9. 

The most widespread of all African Ochna species, 
extending from Yemen, Eritrea and Ethiopia along the 
eastern escarpment to Rhodesia, Botswana, Mozambique 
and Transvaal as far south as Komatipoort; inhabiting arid 
regions, saxicolous or in sandy soil. Flowering in spring. 

Transvaal. — 2229 (Waterpoort): farm Hackthome 608, 
Dongola Reserve, Pole Evans 4436; Wylliespoort, Wer- 
dermann & Oberdieck 1944; Codd 4824; Story 5947 . 2230 
(Messina): Messina, Rogers 19398; Wild 7626. 2231 
(Pafuri): 27 km S.E. of Pafiiri, Mockford 1. 2328 (Balti- 
more): Magoba’s Kloof, Maguire 1468. 2331 (Phalabor- 
wa): Kruger National Park, near Letaba River, Galpin 
15078. 2427 (Thabazimbi): De Hoop trig, beacon. Burn 
Davy sub BOL 18207. 2428 (Nylstroom): Near Nylstroom, 
on lower slopes of Krantzberg, Prosser 1710; Moorddrif, 
Leendertz 2131; Olifantspoort near Nylstroom, Galpin 
13796. 2431 (Acomhoek): 10 km N. of Olifants River, 
Codd 6195. 2529 (Witbank): Loskopdam, Theron 1558; 
1885. 2531 (Komatipoort): Kruger National Park, near 
Kemp’s cottage, Codd 5715; Komati Poort, Pole Evans sub 
PRE 16856. 

10. Ochna glauca Verdoorn in Bothalia 
6: 232 (1951); Robson in F.Z. 2: 239 (1963). 
Type: Transvaal, Soutpansberg District, farm 
De Klundert, Pole Evans 4494 (PRE, holo.). 

Spreading shrubs or small trees. Branches 
greyish-brown, smooth; side branchlets numer- 
ous, short, bearing knob-like spurs which pro- 
duce leaves and flowers; lenticels sparse; young 
parts often covered with wax. Leaves decidu- 



ous, crowded on the apices of the branchlets, 
drooping, often folded, broadly elliptic, c. 3 cm 
long, 1,8 cm broad (in S. African material), 
obtuse at apex and base, midrib prominent, 
margin obscurely serrulate, thin, glaucous, 
metallic green; petiole thin, 4 mm long, usually 
recurved; stipules intrapetiolar, fused. Flowers 
appearing before leaves, solitary, apical, fairly 
small; pedicels 1 cm long, not accrescent in 
fruit but firmer, articulated at the base. Sepals 
elliptic, 5 mm long, green at anthesis, up to 1 
cm long, reddish brown and reflexed in fruit. 
Petals ovate, clawed, 1 1 mm long, yellow. 
Stamens about 40, the filaments c. 4 mm long; 
anthers biporose, c. 2 mm long. Carpels 5, 
attached basally to torus; styles fused, apices 
recurved with slightly swollen stigmas. 
Drupelets oblong-globose, c. 8 mm long, 
brown. Fig. 2: 10. 

Apparently restricted to the northern Transvaal and the 
Matopos in Rhodesia, on granite hills. 

Transvaal. — 2228 (Maasstroom): Kremetartberg area. 
Van Graan & Hardy 483. 2229 (Waterpoort): farm De 
Klundert, Pole Evans 4588; Verdoorn 2292; 2332. 2328 
(Baltimore): Steilloopbrug, Van der Schijff 5330. 

11. Ochna cinnabarina Engl. & Gilg in 
Warb., Kunene-Samb. Exped. 305 (1903); Gilg 
in Bot. Jahrb. 33: 236 (1903); Exell & Men- 
donpa in C.F.A. 1,2: 292 (1951); Robson in 
F.Z. 2: 236 (1963); Schreiber in F.S.W.A. 42: 2 
(1968). Type: Angola, on the Kuito, below 
Longa, on sand hills, Baum 550 (Bf; E, lecto.; 
PRE, photo.). 

Small shrubs up to 1,5 m tall with virgate 
stems bearing short side branches, from a per- 
sistent woody base; lenticels pale, numerous on 
young brown twigs which become greyish and 
rough with age. Leaves deciduous, narrowly 
elliptic, 3-4 cm long and 1-1,5 cm broad, acute 
to obtuse at the apex and base, margin 
spinulose-serrate, leathery, discolorous, midrib 
distinct; stipules intrapetiolar, fused, linear. 
Flowers appearing before the leaves, solitary, 
apical on short lateral spurs; pedicels 1 cm long, 
erect at anthesis, up to 1,5 cm long and spread- 
ing in fruit, articulated below centre. Sepals 
ovate, 1 cm long, obtuse, flat, spreading, 
greenish-yellow, up to 2 cm long and scarlet in 
fruit, forming a loose cup around young fruit, 
spreading when drupelets are ripe. Petals obo- 
vate, clawed, c. 1,5 cm long, bright yellow. 
Stamens c. 50; filaments c. 6 mm long; anthers 
biporose, c. 3 mm long. Carpels 5, attached 

basally; styles fused, terete, apices free, re- 
curved with capitate stigmas. Drupelets 
oblong-globose, c. 1 cm long, black. Fig. 2: 1 1. 

Recorded from Botswana, northern South West Africa, 
southern Angola, Rhodesia and Zambia, in rock fissures, or 
on antheaps, in Kalahari sand. Flowering in November. 

S.W.A. — 1718 (Kuring-kuru): Omuramba Mpungu on 
road from Kuring-kuru to Tsintsabis on dunes of banks, De 
Winter 3873. 1723 (Singalamwe): Singalamwe area, Killick 
& Leistner 3202. 1724 (Katima Mulilo): near Katima 
Mulilo, Pienaar & Vahrmeijer 191. 1819 (Karakuwise): 
Cigarette, N.E. of Karakuwise, Maguire 2271. 1820 

(Tarikora): Masari, Giess 95 1 9. 1821 (Andara): 23 km W. 
of Andara Mission Station on road to Nyangana, De Winter 
& Wiss 4225. 

12. Ochna barbosa t Robson in Bol. Soc. 
Brot., Ser. 2, 36: 18 (1962); in F.Z. 2: 238 
(1963). Type: Mozambique, Vila Luis, Bar- 
bosa & Lemos sub Barbosa 7895 (LISC, holo.; 
COI; K; LMJ). 

A slender tree up to 8 m tall or a shrub with 
light brown, slightly rough bark, not flaking; 
branchlets with the epidermis peeling off in 
narrow membranous strips; lenticels many, 
scattered; woody ovoid galls often present. 
Leaves shed before flowering or shortly after, 
narrowly elliptic to elliptic, 3-5 cm long and 
1,2- 1,5 cm broad (in S. African material seen), 
apex rounded, base acute, margin indistinctly 
serrulate (appearing smooth), coriaceous, with 
distinct reticulate venation; petiole 1 mm; 
stipules intrapetiolar, fused, entire. Flowers 
scented, terminating short spurs, solitary; 
pedicels c. 4 mm long at anthesis, up to 1 (-2) 
cm in fruit, erect, articulated at the base. Sepals 
about 9 mm long at anthesis, about 2 cm long 
and pink or red in fruit, enclosing young 
drupelets, spreading when fruit is ripe. Petals 
yellow, obovate-cuneate, c. 12 mm long and 8 
mm broad. Stamens c. 50, filaments c. 2,5 mm; 
anthers 1,5 mm, biporose. Carpels 5, attached 
basally to receptacle; styles fused, terete, with 
free apical, recurved stigmatic tips or com- 
pletely fused with stigma apically, discoid, 
5-lobed. Drupelets oblong-globose, 1 cm long, 
black. Fig. 2: 12. 

Occurs in Natal, southern Mozambique, and south- 
eastern Rhodesia. A sand-loving tree or shrub inhabiting 
coastal dune forest, locally common where found; extend- 
ing inland along low lying areas to south-eastern Rhodesia. 

Natal. — 2632 (Bela Vista): 7 km W. of Muzi, Moll & 
Nel 5590. 2732 (Ubombo): Sihangwane Store, Moll 4410; 
Lake Sibayi, Venter 5818; Mbazwane-Mseleni Road, 
Pooley 1512. 3030 (Port Shepstone): Isipingo Beach, Ward 
1239; 3759; 5887. 



Hybrids: Moll & Strey 3928 (2632 Bela Vista), near 
Banga Nek, and Ward 3611 (2732 Ubombo), Mkuze Game 
Reserve, both collected in dune forest, northern Natal, ap- 
pear to be hybrids between O. barbosae and O. natalitia. 


Brackenridgea A. Gray, Bot. U.S. Expl. Exp. 
Bot. 1: 361, t. 42 (1854); Walp., Ann. 4: 421 
(1857); Gilg in Pflanzenfam. ed. 2,21: 74 
(1925); Robson in F.Z. 2: 252 (1963); Schreiber 
in F.S.W.A. 42: 1 (1968). Type species: B. 
nitida A. Gray from Fiji. 

Trees, shrubs or soboliferous dwarf 
shrubs, glabrous, forming a yellow pigment 
under the bark. Leaves simple, shortly petio- 
late, with the primary lateral nerves curving 
upwards, almost parallel to the margin, the 
margin closely serrulate with the teeth gland- 
tipped; petioles short; stipules deeply divided 
into subulate segments, striate, persistent at 
least on first year shoots in African species. 
Flowers variously arranged, usually in axillary 
fascicles or reduced to 1 flower; bracts resemb- 
ling stipules. Sepals (4) 5, imbricate, white or 
pink, turning red and accrescent in fruit. Petals 
(4) 5, white or pale pink, fugaceous. Stamens 
10-20, free; filaments persistent; anthers bilocu- 
lar, basifixed, opening longitudinally, caducous. 
Carpels 5-10, attached basally to torus; style 
entire; stigma capitate. Drupelets globose, 
black attached basally to the red swollen torus; 
seeds without endosperm, embryo curved. 
Fig. 3. 

Species about 12, widely distributed in S.E. Asia and 
Australia; 4 in tropical and subtropical Africa and Madagas- 

car. One tropical to subtropical species recorded from South 
West Africa, usually in sandy soil. 

Brackenridgea arenaria (De Wild. & 
Dur.) Robson in Bol. Soc. Brot., Ser. 2, 36: 37 
(1962); in F.Z. 2: 254, t. 46, f.b (1963); 
Schreiber in F.S.W.A. 42: 2 (1968). Type: 
Zaire, Kisantu, Gillett 68 (BR, holo.). 

Ochna arenaria De Wild. & Dur. in Bull. Herb. Boiss. 2, 
1: 7 (1900). Type as above. 

Small rhizomatous shrublets with virgate, 
grooved branches, up to 1 m tall (rarely some- 
what taller); wood yellow below grey-brown 
peeling bark. Leaves deciduous, erect, close 
together, alternate, narrowly obovate, c. 8 cm 
long, 2 cm broad, usually obtuse above, apicu- 
late, attenuate towards the base, forming a short 
petiole, margin closely and shallowly serrulate 
with the teeth bearing ovoid glands, firm, shiny, 
the lateral veins curved upwards along margin; 
stipules deeply dissected into subulate seg- 
ments, striate, intrapetiolar, persisting on first 
year shoots. Flowers in short, axillary, sessile 
(1) 3-6-flowered fascicles; the bracts resemb- 
ling the stipules; pedicels c. 8 mm long at 
anthesis, somewhat longer in fruit, articulated at 
the base. Sepals narrowly oblong, c. 5 mm at 
anthesis, up to 1 cm in fruit, pink or red, 
spreading or reflexed. Petals narrowly obovate, 
somewhat longer than sepals, white or pinkish, 
reflexed at anthesis. Stamens about 20; anthers 
opening by longitudinal slits. Carpels 5, 
rounded, inserted basally on the torus; style 
terete, stigma apical, small. Fig. 3. 

Widespread in tropical and southern subtropical reg- 
ions, as far south as northern South West Africa; in sandy 

S.W.A. — 1723 (Singalamwe): E. of base E.C.Z., 
Pienaar & Vahrmeijer 207. 1821 (Andara): Caprivi Strip, 
between Bagani Camp and Mahango, De Winter & Wiss 
4395; 16 km from Bagani towards Angola border, Watt 2. 



Fig. 3. — 1, Brackenridgea arenaria, leafy branch, x 1 (Pienaar & Vahrmeijer 207); a, leaf-margin showing glands, X 4; 
b, base of leaf showing stipules, X 4; c, portion of flowering twig, x 1, ( Pocock 1925); d, anther-locules with 
longitudinal dehiscence, x 10. 




(nom. alt. Guttiferae) 

by D. J. B. Killick and N. K. B. Robson* 

Trees, shrubs or herbs, with resinous juice. Leaves opposite, sub-opposite or whorled, simple, 
exstipulate, glandular. Flowers actinomorphic, bisexual, dioecious or polygamous. Sepals 2-6, 
quincuncial or decussate. Petals same number as sepals, free, usually convolute, sometimes 
imbricate or decussate, alternating with sepals. Androecium basically of two whorls of stamen 
fascicles, the outer often sterile (fasciclodes) or absent, filaments variously united or free, inner 
fascicles variously united or free, very rarely each one reduced to a single stamen. Ovary superior, 
1-many-locular, placentation usually axile, sometimes ± parietal, ovules 1-many; styles free, ± 
united or absent; stigmas equal in number to loculi. Fruit a septicidal (rarely loculicidal) capsule or 
berry or drupe. Seeds sometimes winged, carinate or arillate, without endosperm. 

Almost exclusively tropical in distribution and composed of about 40 genera and 1000 species (Willis, 1966). 2 genera 
and 8 species occur in our area. 

Flowers bisexual; sepals 5; styles (2) 3-5, elongate; ovules several-many per 

loculus 1. Hypericum 

Flowers polygamous or dioecious, rarely bisexual; sepals 4; styles absent and stigmas sessile; 

ovules 1 per loculus 2. Garcinia 

5168 1. HYPERICUM 

Hypericum L., Sp. PI. 1: 443 (1753); Gen. PI. ed. 5: 341 (1754); DC., Prodr. 1: 550 (1823); Sond. 
in F.C. 1: 117 (1860); Benth. & Hook.f., Gen. PI. 1,1: 165 (1862); Oliv. in F.T.A. 1: 154 (1868); 
Keller in Pflanzenfam. ed. 2, 21: 175 (1929); Bredell in Bothalia 3: 574 (1939); Phill. , Gen. ed. 2: 
506 (1951); Milne-Redh. in F.T.E.A. Hypericaceae: 1 (1953); Robson in F.Z. 1: 379 (1961). Type 
species: H. perforatum L. 

Trees, shrubs or herbs. Leaves opposite, sessile, amplexicaul or shortly petiolate, entire, with 
translucent glandular dots or linear striae and frequently with dark submarginal glandular dots. 
Flowers terminal, solitary or cymose, sometimes forming a corymb or panicle, bisexual. Sepals 
usually 5, equal, subequal or unequal, with translucent gland dots or striae, with or without dark 
often submarginal dots and sometimes with stalked marginal glands. Petals usually 5, usually 
yellow, often red-tinged, asymmetrical, with translucent or dark glandular dots or striae. Stamens 
usually very numerous, free or connate at the base into 3-5 distinct or indistinct groups; filaments 
thread-like; anthers with or without yellow or black gland. Ovary superior, sessile, ovoid or 
subround, 3-5-locular, with few to many ovules on parietal placentas in each loculus; styles 3-5, 
free or variously united; stigmas small, capitate. Fruit a septicidal capsule with few to many seeds, 
rarely indehiscent. Seeds minute, cylindric to ovoid, straight or slightly curved; testa thin ribbed, 
punctate or smooth; embryo straight or curved. 

♦Department of Botany, British Museum (Natural History), Cromwell Road, London SW7 5BD. 



A large genus of about 400 species, widely dispersed, especially in the northern hemisphere. Six species in Southern 

The generic name is derived from the Greek name for St. John’s Wort, viz. hypereikon, i.e. hyper + eikon = above + 
image. The Greeks used plants of this genus to decorate religious images in order to ward off evil spirits, especially around 
midsummer’s eve (St. John’s eve). 

Several species of Hypericum are cultivated as ornamentals in Southern Africa, namely H. calycinum L., H. 
monogyrum L. and H kalmianum L. H. perforatum L. is a troublesome weed in the Stellenbosch area of the south-western 
Cape. Species of Hypericum contain the active principle hypericin, which renders the plants poisonous to stock through 


Styles 5, wholly or partially united; shrubs: 

Flowers solitary at ends of branches; leaves 0,8-2,3 cm long 1. H. revolutum 

Flowers in few-to many-flowered terminal corymbose cymes; leaves 4—8,5 cm long 2. H. roeperanum 

Styles 3-5, free; herbs: 

Stems quadrangular; sepals and petals without dark dots 3. H. lalandii 

Stems terete; sepals usually with dark dots: 

Sepals acute, subequal 4. H. aethiopicum 

Sepals obtuse to rounded, unequal: 

Stems erect, 20-45 cm high; leaves sessile, rarely shortly petiolate 5. H. natalense 

Stems procumbent, 6-10 cm high; leaves shortly petiolate 6. H. wilmsii 

1. Hypericum revolutum Vahl, Symb. 
Bot. 1: 66 (1790); Christensen in Dansk Bot. 
Ark. 4,3: 39 (1922); Robson in Kew Bull. 14: 
251 (1960); in F.Z. 1: 381 (1961); Moggi & 
Pisacchi in Webbia 22: 236, 1. 1 (1967). Lec- 
totype: “Arabia felix, in montibus”, Forskal 
796 (C!). 

H. kalmii Forsk., FI. Aegypt.-Arab. CXV1II (1775), 
nomen nudum. H. lanceolatum Lam., Encycl. 4: 145 
(1797); Oliv. in F.T.A. 1: 156 (1868); Bak.f. in J. Linn. 
Soc. (Bot.) 40: 26 (191 1); Eyles in Trans. R. Soc. S. Afr. 5: 
420 (1916); Good in J. Bot., Lond. 65: 330, t.582, fig. 1 
(1927); Staner in Bull. Jard. Bot. Brux. 13: 74 (1934); 
Robyns, FI. Parc Nat. Alb. 1: 620, t.62 (1948); Perrier in FI. 
Madag. Hypericaceae: 3 (1951); Milne-Redh. in F.T.E.A. 
Hypericaceae: 4 (1953); in Mem. N.Y. Bot. Gard. 8: 221 
(1953); Keay & Milne-Redh. in F.W.T.A. ed. 2,1: 287, 
1. 109 (1954); Robson in Kew Bull. 12: 444 (1957). Type: 
Reunion (“Bourbon”), Commerson (LINN-SM, iso.!). H. 
leucoptychodes Steud. ex A. Rich., Tent. FI. Abyss. 1: 96 
(1847); Good, l.c. t.582, fig. 3-5 (1927); Norlindh in Bot. 
Notiser 1934: 100 (1934); Bredell in Bothalia 3: 580 (1939); 
Phill. in Flow. PI. S. Afr. 20: t.787 (1940); Brenan, 
Checklist Tang. Terr. 249 (1949). Syntypes: Ethiopia, near 
Dschenausa, Schimper 834 (K!); Mt. Bachit, Schimper 
1177 (K!). H. lanuriense De Wild., PI. Bequaert. 5: 403 
(1932); Robyns, l.c. 622 (1948). Type: Zaire, Ruwenzori, 
Ruaroli (Lanuri) Valley, Bequaert 4460 (BR, holo.; K, 

Shrub up to 3 m high (12 m in tropical 
Africa). Stems 4-angled when young, but be- 
coming ± terete, often with scaly bark. Leaves 
sessile, narrowly elliptic, 0,8-2, 5 cm long, 3-6 
mm wide, apex acute, base cuneate, narrowing 
to a clasping base, venation sparsely pinnate, 
tertiary venation variously conspicuous, with 
translucent longitudinal glands or striae and few 
dark submarginal glands. Flowers solitary, 

terminal. Sepals unequal, ovate, 6-10 mm long, 

3- 5 mm wide, concave, with few dark submar- 
ginal glands and often numerous stalked mar- 
ginal glands. Petals obovate-spathulate, 2-3 cm 
long, 0,8- 1,8 cm wide, yellow, without dark 
submarginal gland dots. Stamens in 5 bundles 
of 20-30 each; filaments 8-15 mm long; anthers 
c. 1-1,3 mm long, with apical gland. Ovary 
ovoid, 4—6 mm long, 3-5 mm wide; styles 5, 

4— 8 mm long, united except for upper 1,5-4 
mm. Capsule ovoid, 1-1,2 cm long, 0,7-1, 1 cm 
wide, 5-valved. 

Found on streambanks and the edge of forest in the 
Transvaal, Natal, Swaziland and the eastern Cape. Also 
occurring in tropical Africa, Fernando Po, Malagasy, the 
Comoro Islands and Reunion. 

Transvaal. — 2230 (Messina): Entabeni Forest Station, 
Codd 4206. 2329 (Pietersburg): Houtboschdorp, Van Vuu- 
ren 1203. 2330 (Tzaneen): Westfalia Estates, Duiwelskloof, 
Scheepers 412. 2430 (Pilgrims Rest): Reitz’s Grave, 
Mariepskop, Van der Schijff 6803. 2530 (Lydenburg): 
Duiwelskantoor, Thode A 1565. 2627 (Potchefstroom): 
Witpoortjie, near Krugersdorp, Grant 2687. 

Swaziland. — 2531 (Komatipoort): near Havelock, 

Compton 30123. 2631 (Mbabane): Mbabane, Rogers 
11483; Miller SI 122. 

Natal. — 2831 (Nkandla): Nkandla Forest, Pentz 351; 
Nhlazatshe, Ward 3410. 2930 (Pietermaritzburg): Illovo 
River, near Richmond, Wood 1995 (NH); Pateni Estates, 
Strey 7636. 

Cape. — 3029 (Kokstad): Malowe, Tyson 3044. 

An attractive plant with large, yellow flowers, known 
as the Forest Primrose or Curry Bush, because of the 
curry-like odour given off by the leaves, particularly when 
crushed. According to Watt & Brandwijk (1962) the plant 
has been used as a source of balsam. In the Haenertsburg 
area of the Transvaal it is called “Nature’s Fire Break” 
because, situated as it often is at the edge of forests, it is said 



to prevent fire from penetrating forest ( Hodgson s.n. in 
PRE). However, it is doubtful whether the plant is particu- 
larly fire-resistant. Experimentally it has induced non-icteric 
photosensitization in sheep, due apparently to the presence 
of hypericin (Watt & Brandwijk, 1962). 

2. Hypericum roeperanum Schimp. ex 
A. Rich., Tent. FI. Abyss. 1: 96 (1847); Keller 
in Pflanzenfam. ed. 2, 21: 176 (1925); Good in 
J. Bot., Lond. 65: 331, t.582, fig. 6 (1927); 
Staner in Bull. Jard. Bot. Brux. 13: 76 (1934); 
Exell & Mendonpa in C.F.A. 1,1: 119 (1937); 
Bredell in Bothalia 3: 582 (1939); Brenan, 
Checklist Tang. Terr. 250 (1949); Suesseng. & 
Merxm. in Proc. & Trans. Rhod. Sci. Ass. 43: 
88 (1951); Milne-Redh. in F.T.E.A. 

Hypericaceae:3 (1953); Robson in Kew Bull. 
12: 444 (1957); in F.Z. 1: 380 (1961); Moggi & 
Pisacchi in Webbia 22: 244, t.2 (1967). Type: 
Ethiopia, “crescit ad montem Kubbi, prope 
Adoua et ad montem Aber prope Dschenausa in 
provincia Semiene”, Schimper 866 (P, lecto.; 
BM!; FI; G; K!; LE; MO). 

var. roeperanum. 

Moggi & Pisacchi in Webbia 22: 247 

H. quartinianum A. Rich. var. roeperanum (Schimp. ex 
A. Rich.) Engl., Hochgebirgsfl. Trop. Afr. 307 (1892). 

Shrub or small tree, up to 5 m high. Stems 
± terete, smooth, glabrous. Leaves sessile, 
elliptic, 4—8,5 cm long, 1,2-3, 4 cm wide, apex 
acute or obtuse (sometimes apiculate when 
obtuse), base cuneate, clasping, with several 
pairs of secondary veins and conspicuous re- 
ticulation of tertiary veins each reticulation 
containing a translucent glandular dot, dark 
glands on margin. Flowers in few- to many- 
flowered terminal corymbose cymes. Sepals 
unequal, ovate, 4—6 mm long, 3,5-4 mm wide, 
acute-obtuse, with submarginal dark gland dots 
and marginal stalked glands. Petals obovate- 
spathulate, 2-3 cm long, 1-1,2 cm wide, bright 
yellow, with few or no submarginal gland dots. 
Stamens in 5 bundles of about 45 each; fila- 
ments 1,2- 1,4 cm long; anthers c. 0,75 mm 
long, with apical gland. Ovary ovoid, 5-7 mm 
long; styles 5, united to apex, 8-10 mm long. 
Capsule ovoid, 1-1,8 cm long, 0,6-1 cm wide, 

Widely distributed in Africa from Sudan and Ethiopia 
southwards to the north-eastern Transvaal and Mozam- 
bique. Apparently restricted in South Africa to the 
Mariepskop area. Usually occurring on streambanks, in 
woodland or at the edge of forest. 

Transvaal. — 2430 (Pilgrims Rest): Mariepskop area, 
Killick <£ Strey 2424; Van der Schijff 4975; Blyde River 
Road, Van der Schijff 5529. 

Differing from H. revolutum Vahl in that the flowers 
are in few- to many-flowered terminal corymbose cymes 
instead of solitary and in the larger leaves with densely 
reticulate venation. Var. schimperi (Hochst. ex A. Rich.) 
Moggi & Pisacchi is restricted to Ethiopia. 

3. Hypericum lalandii Choisy in DC., 
Prodr. 1: 550 (1824); Oliv. in F.T.A. 1: 155 
(1868); Gibbs in J. Linn. Soc. (Bot.) 37: 430 
(1906); Bak.f. in J. Linn. Soc. (Bot.) 40: 26 
(1911); Eyles in Trans. R. Soc. S. Afr. 5: 420 
(1916); Keller in Bot. Jahrb. 58: 197 (1923); in 
Pflanzenfam. ed. 2, 21: 181 (1925); Burtt Davy, 
FI. Transv. 1: 251 (1926); Staner in Bull. Jard. 
Bot. Brux. 13: 70 (1934); Norlindh in Bot. 
Notiser 1934: 102 (1934); Gomes e Sousa in 
Bol. Soc. Estud. Col. Mocamb. 26: 42 (1935); 
Exell & Mendonpa in C.F.A. 1,1: 120 (1937); 
Bredell in Bothalia 3: 575 (1939); Adamson in 
FI. Cape Penins. 588 (1950); Suesseng. & 
Merxm. in Proc. & Trans. Rhod. Sci. Ass. 43: 
88 (1951); Perrier in FI. Madag. Hypericaceae: 
14, 1. 1, figs. 7 & 8 (1951); Milne-Redh. in 
F.T.E.A. Hypericaceae: 7 (1953); Robson in 
F.Z. 1: 385 (1961); in Garcia de Orta 1: 85 
(1973). Type: Cape, Laland s.n. (P, holo.; PRE, 

H. lalandii Choisy var. lanceolata Sond. in F.C. 1: 118 
(1860). Type: Port Natal, Drege d (S !). — var. latifolia 
Sond., l.c. Type: Uitenhage, Zeyher s.n. (S !). — var. 
macropetala Sond., l.c. Type: Drege c (S, holo.!; BM!; 
K!). — var. lanceolatum Keller in Bull. Herb. Boiss. Ser. 
2,8: 187 (1908). Syntypes: Natal, Gueinzius s.n. (Bf); 
Gerrard 190 (K!; PRE, photo.; W!). — var. transvaalense 
Bredell in Bothalia 3: 577 (1939). Type: Transvaal, 
Rehmann 6608 (PRE, lecto.!; BM!). 

Herb, 4—40 cm high. Stems from a woody 
underground rootstock, erect or decumbent, 
rarely procumbent, branched or unbranched, 
4-ribbed, glabrous, glandular or eglandular. 
Leaves sessile, ascending; blade very variable 
in shape and size, upper and middle leaves 
usually narrowly ovate, narrowly elliptic or 
linear, basal leaves broader and shorter, 3-36 
mm long, 0,5-6 mm wide, apex acute or 
obtuse* base clasping, venation not con- 
spicuous, pellucid-glandular punctate. Flowers 
up to c. 50 in a loose dichasial cyme or solitary. 
Sepals ± equal, narrowly ovate, 5,5-7 mm 
long, 1,2- 1,5 mm wide, long-acute, with trans- 
lucent longitudinal veins and gland dots. Petals 
1-2 times as long as sepals, yellow or orange, 
sometimes marked with red, distinctly veined. 



Stamens 40-60, irregularly arranged; filaments 
4—6 mm long; anthers c. 0,3 mm long. Ovary 
ovoid, 1,5-3, 5 mm long, 1-1,5 mm wide, 
1 -locular; styles 3, free, 1,5-2 mm long; stig- 
mas capitate. Fruit capsular, (2) 3-4-valved. 

Widely distributed in Africa from Nigeria and the 
Sudan southwards to the Cape Province. Also in Madagas- 

S.W.A. — 2017 (Waterberg): Waterberg Plateau, Boss 
sub TRV 34997; Okosongomingo, Dinter 1758 (SAM). 

Transvaal. — 2230 (Messina): Entabeni, Obermeyer 
sub TRV 30071. 2327 (Ellisras): Ellisras, Bayliss 2393 
(NBG). 2328 (Baltimore): on Mohlakeng Plateau, Blauw- 
berg, Codd & Dyer 8992. 2329 (Pietersburg): Eloutbosch, 
Rehmann 6340. 2330 (Tzaneen): Westfalia, Scheepers 
1093; 1116. 2429 (Nylstroom): Mosdene Farm, 

Naboomspruit, Germishuysen 49. 2430 (Pilgrims Rest): 
Mariepskop, Van der Schijff 6021 . 2526 (Zeerust): Zwart- 
ruggens, Fouche 12. 2527 (Rustenburg): Rustenburg Na- 
ture Reserve, Jacobsen 804. 2528 (Pretoria): Wonderboom, 
Smith 2291. 2529 (Witbank): Loskop Dam — Donkerhoek, 
Theron 1603. 2530 (Lydenburg): Rosehaugh, Mogg 13962. 
2531 (Komatipoort): Numbi, Van der Schijff 4158. 2627 
(Potchefstroom): Buffelshoek, Potchefstroom, Louw 1527. 
2628 (Johannesburg): Heidelberg, Leendertz 1030. 2629 
(Bethal): Nooitgedacht, Ermelo, Henrici 1348. 2630 
(Carolina): Bereton Park Outspan, near Sheepmoor, De 
Winter 7520. 2729 (Volksrust): Amersfoort, Sidey 3504. 
2730 (Vryheid): Oshoek, Wakkerstroom, Devenish 1094. 

O.F.S.— 2727 (Kroonstad): Bothaville, Goossens 1216. 
2827 (Senekal): Willem Pretorius Game Reserve, Leistner 
2969. 2828 (Bethlehem): Bethlehem, Phillips 3215. 2829 
(Harrismith): Harrismith, Sankey 21 (K). 

Swaziland. — 2631 (Mbabane): 5 km N.E. of Forbes’s 
Reef, Schlieben 9531; Poliniane River, Compton 26491; 
Nduma, Compton 27269. 

Natal. — 2632 (Bela Vista): Maputa Store, Moll & Strey 
3875. 2729 (Volksrust): “Glen Atholl”, near Charlestown 
& Volksrust, Smith 5697. 2730 (Vryheid): Tweekloof 
(Altemooi), Thode A 1145. 2731 (Louwsburg): Nongoma 
Townlands, Gerstner 4612. 2732 (Ubombo): Sibayi, Vahr- 
meijer 554. 2828 (Bethlehem): Royal Natal National Park, 
Galpin 10177. 2829 (Harrismith): Cathedral Peak Forest 
Station, Killick 1173. 2830 (Dundee): Qudeni, Gerstner 
1950. 2831 (Nkandla): Melmoth, Mogg 6052. 2832 
(Mtubatuba): St. Lucia Park, Dutton 35. 2929 (Underberg): 
between Loteni and Giants Castle, Killick 3915. 2930 
(Pietermaritzburg): “Ehlatini Farm”, Karkloof, Moll 3460. 
2931 (Stanger): Wentworth, Durban, Ward 6463. 3029 
(Kokstad): Weza, Ingeli, Strey 6316. 3030 (Port 

Shepstone): Uvongo, Strey 4889. 

Lesotho. — 2828 (Bethlehem): Leribe, Dieterlen 678. 
2927 (Maseru): Mazenod, Jacot Guillarmod 805. 2929 
(Underberg): Sehlabathebe, Jacot Guillarmod, Getliffe & 
Mzamane 86. 

Cape. — 2822 (Glen Lyon): Witsand, Hay Division, 
Acocks 2173. 3029 (Kokstad): Clydesdale, Tyson 2097. 
3128 (Umtata): Baziya Forest Station, Mauve 4862. 3129 
(Port St. Johns): Ndindini, Strey 10109. 3227 (Stutterheim): 
Evelyn Valley, Leighton 26 72. 3228 (Butterworth): Ken- 
tani, Pegler 117, pro parte. 3322 (Oudtshoom): George, 
Marloth 2545. 3225 (Port Elizabeth): 1 1 km from Ann’s 
Villa on Zuurberg Pass, Acocks 20298. 3326 

(Grahamstown): near Dassie Krantz, Britten 627. 3327 
(East London): near East London, Acocks 21848. 3418 

(Simonstown): Sir Lowry’s Pass, Marloth 4866. 3420 
(Bredasdorp): Swellendam, Compton 584 (NBG). 3422 
(Mossel Bay): Vrijers Berg, Muir 2043. 3423 (Knysna): 
Harkerville, Breyer sub TRV 23914. 

An extremely variable species in habit, leaf shape and 
size, flower size and colour and density of punctate glands. 

4. Hypericum aethiopicum Thunb., 
Prodr. 2: 138 (1800). Types: Cape Province, 
Thunberg a (UPS, lecto. !); Thunberg (3 (UPS!). 

Perennial herb, 10-30 cm high. Stems 
erect or decumbent, unbranched, arising from 
an underground crown, terete or sometimes ± 
flattened or 2-lined above, eglandular or spotted 
with dark glands. Leaves sessile, ovate to 
broadly ovate, 0, 8-2,3 cm long, 0,4— 1,5 cm 
wide, apex obtuse to rounded, base cordate- 
amplexicaul, margins often revolute, with 
numerous translucent punctate glands and dark 
marginal glands. Flowers in lax or compact 
terminal few- to many-flowered cymes, rarely 
solitary; pedicels 2-4 mm long, except in fruit. 
Sepals subequal, ovate, 5-8 mm long, 1,5-2 
mm wide, apex sharply acute or acuminate, 
with translucent gland dots and striae and with 
or without dark gland dots both marginal and on 
lamina, with or without stalked marginal 
glands. Petals 10-13 mm long, 2-4 times as 
long as sepals, primrose yellow, usually red- 
tinged, with dark gland dots and translucent 
gland dots or striae, distinctly veined. Stamens 
c. 50-70, irregularly arranged or in 3 or 4 
indistinct groups, with filaments united at ex- 
treme base; anthers 0,4— 0,5 mm long, often 
with black or orange glands at apex of connec- 
tive. Ovary ovoid, 2,5-4 mm long, 1,5-2 mm 
wide, 3 (4)-locular; styles 3, free, 4,5-6,5 mm 
long; stigmas ± capitate. Fruit capsular, ovoid, 
6 mm long, 3 mm wide, 3-4-valved, erect. 

A grassland herb occurring in Rhodesia, Mozambique, 
Angola, Lesotho, Swaziland and South Africa. 

Two subspecies are recognized and may be keyed out 
as follows: 

Stems eglandular, sepals and bracts with marginal 

stalked glands (a) subsp. aethiopicum 

Stems glandular, sepals and bracts without marginal 

stalked glands (b) subsp. sonderi 

(a) subsp. aethiopicum. 

Robson in Kew Bull. 12: 439 (1957); in F.Z. 1: 
382 (1961). 

H. aethiopicum Thunb., Prodr. 2: 138 (1800); Sond. in 
F.C. 1: 117 (1860), pro parte. 


Fig. 4. — 1, Hypericum aethiopicum subsp. aethiopicum, portion of plant, X 1; la, sepal, x 7; lb, petal, x 7 (Flanagan 
2142). 2, H. aethiopicum subsp. sonderi, portion of plant, x 1; 2a, sepal, X 7; 2b, petal, X 7 (Hafstrom Acocks 



This subspecies is confined to the Cape Province 
between the Korinte River near Riversdale and the moun- 
tains of Griqualand East. Fig. 4: I. 

Cape. — 3029 (Kokstad): mountain side, Griqualand East, 
Tyson 1376, pro parte. 3321 (Ladismith): Korinte River, 
Riversdale, Muir 855. 3226 (Fort Beaufort): Hogsback 
Forest Reserve, Dahlstrand 1810. 3227 (Stutterheim): Stut- 
terheim Commonage, Acocks 9544. 3322 (Oudtshoom): 
Kamanassie Hills near George, Prior s.n. (K). 3325 (Port 
Elizabeth): Zuurberg Sanatorium, Long 225 (K). 3326 
(Grahamstown): Kolsrand, Johnson 1085; Howiesons 
Poort, MacOwan 397 (BM; K). 3423 (Knysna): Knysna, 
Schonland 3498. 

(b) subsp. sonderi (Bredell) Robson in 
Kew Bull. 12: 440 (1957); in F.Z. 1: 382 
(1961). Type: Natal, near Camperdown, 
Schlechter 3270 (K, lecto. !; G!; PRE!). 

H. aethiopicum Thunb. var. glaucescens Sond. in F.C. 1: 
118 (1860); Burtt Davy in FI. Transv. 1: 251 (1926). 
Syntypes: Transvaal, Apies River and Magaliesberg, 
Zeyher s.n. (Burke s.n. from Magaliesberg in BM! and K! 
may be type material — also Zeyher 149 in BM!). H. 
aethiopicum sensu Sond., l.c., pro parte; sensu Bak.f. in J. 
Linn. Soc. (Bot.) 40: 26 (1911); Eyles in Trans. R. Soc. S. 
Afr. 5: 420 (1916); Norlindh in Bot. Notiser 1934: 101 
(1934); Exell & Mendonpa in C.F.A. 1,1: 120 (1937). H. 
sonderi Bredell in Bothalia 3: 578 (1939); Verdoom in 
Flow. PI. S. Afr. 23: t. 897 (1943). — var. transvaalense 
Bredell, l.c. 579. Syntypes: Transvaal, Woodbush, Mogg 
13996 (PRE!); Shiluvane, Junod 4290 (PRE!); without 
precise locality. Wager sub TRV 7223 (PRE!), pro parte. 

Found in the Transvaal, Orange Free State, Swaziland, 
Natal, Lesotho, the eastern Cape and south tropical Africa. 
Fig. 4: 2. 

Transvaal. — 2230 (Messina): near Entabeni, Hutchin- 
son & Gillett 4238 (BM; K). 2330 (Tzaneen): Pigeon Hole, 
McCullum 17; Woodbush, Mogg 14711. 2427 

(Thabazimbi): below Waterberg, Hutchinson 2291. 2430 
(Pilgrims Rest): The Downs, Junod 4290. 2527 (Rusten- 
burg): 2,8 km N.E. of Derby Station, Acocks 21971 . 2528 
(Pretoria): Apies River, Smith 207. 2530 (Lydenburg): 
Rosehaugh, Mogg 13960. 2531 (Komatipoort): Saddleback 
Mountain, Barberton, Galpin 1028. 2626 (Klerksdorp): 
Goedgedacht, Ventersdorp District, Sutton 502. 2627 
(Potchefstroom): “Uitkomst”, near Skeerpoort, Codd 
10097. 2628 (Johannesburg): Johannesburg, Bredenkamp 
742. 2629 (Bethal): near Amsterdam, Van derMerwe 1260. 
2730 (Vryheid): Piet Retief, Galpin 9638. 

O.F.S. — 2828 (Bethlehem): “Wyndford”, Fouriesburg, 
Gemmell sub BLFU 5812. 2829 (Harrismith): “Grootvlei”, 
near Swinburne, Jacobsz 33. 2927 (Maseru): Ladybrand, 
Werdermann & Oberdieck 1566. 

Swaziland. — 2531 (Komatipoort): Havelock Mine, 
Miller 3074. 2631 (Mbabane): Verdun, Hlatikulu, Compton 

Natal. — 2730 (Vryheid): Tweekloof (Altemooi), Thode 
A1146. 2731 (Louwsburg): Nongoma Townlands, Gerstner 
4611 . 2828 (Bethlehem): Royal Natal National Park, Codd 
2784. 2829 (Harrismith): Cathedral Peak, Goodier 350. 
2830 (Dundee): hill above Dundee Reservoir, Edwards 
1088. 2831 (Nkandla): Empangeni Village, Venter 3251. 
2832 (Mtubatuba): Ezinkakeni, Hluhluwe Game Reserve, 
Hitchens 745. 2929 (Underberg): Cathkin Park, Howlett 94. 

2930 (Pietermaritzburg): Balgowan, Mogg 3516. 3030 
(Port Shepstone): Ifafa, Rudatis 246 (BM; K); Marina 
Beach, Port Shepstone, Slrey 5950. 

Lesotho. — 2828 (Bethlehem): 5 km S. of Buthe-Buthe, 
Galpin s.n. 2927 (Maseru): bank of the Catai River, 
Mafeteng Distr., Dieterlen 1293. 

Cape. — 3029 (Kokstad): 1 km from Bizana on Kokstad 
road. Story 597. 

There is a certain amount of overlap between subsp. 
aethiopicum and subsp. sonderi: occasionally the odd 
specimen of subsp. aethiopicum has glandular stems, while 
specimens of subsp. sonderi sometimes have eglandular or 
nearly eglandular stems. According to Robson (l.c. 439), 
some specimens of subsp. sonderi from the Melsetter region 
of Rhodesia have sepals bearing stalked glands. This 
incomplete correlation between stem and sepal characters 
led Robson to treat these two taxa as subspecies rather than 

According to a note on Wood 225 in Herb. Kew, the 
leaves of the plant are used to make a lotion for inflamed or 
diseased eyes; and the Zulu name is Insukubile, which 
means “two days”. 

5. Hypericum natalense Wood & Evans 
in J. Bot. 35: 487 (1897); Burtt Davy, FI. 
Transv. 1: 25 (1926); Keller in Pflanzenfam. 21: 
177 (1929); Bredell in Bothalia 3: 579 (1939); 
Robson in Kew Bull. 12: 440 (1957). Type: 
Natal, near bank of Mooi River, 1 200-1 500 
m. Wood 4034 (NH, holo.!; BM!; BOL!; G!; 
K!; PRE, photo.). 

H. woodii Keller in Bot. Jahrb. 58: 193 (1923), nom. 
superfl. Type: as for H. natalense Wood & Evans. H. 
natalense Wood & Evans var. petiolatum Bredell in 
Bothalia 3: 580 (1939). Type: Natal, Camperdown District, 
Franks sub NH 12968 (NH, holo.!; PRE, photo.). 

Herb up to 45 cm high. Stems 1-many, 
from a persistent underground rootstock, erect, 
much-branched, round and smooth, glabrous. 
Leaves sessile or rarely shortly petiolate, ellip- 
tic, broadly elliptic or obovate, 0,9- 1,9 cm 
long, 0,5- 1,2 cm wide, apex obtuse to rounded, 
3-7-nerved, tertiary venation fairly conspicu- 
ous, punctate with submarginal dark dots. 
Flowers terminal with flowering branches from 
uppermost pair of nodes. Sepals unequal, ellip- 
tic, obovate or spathulate, 4,5-8 mm long, 
1,5-4 mm wide, apex rounded, slightly apicu- 
late, distinctly veined, densely yellow-punctate 
with few dark submarginal gland dots. Petals 
elliptic, oblong or spathulate, 6,5-7 mm long, 
1,7-2, 5 mm wide, yellow, with 0-few yellow 
gland dots and occasional dark dots near apex. 
Stamens indefinite, 3-5 mm long; anthers 0,3 
mm long, each with a dark gland at end of 
connective. Ovary ovoid, 2,5-3 mm long, 2 
mm wide; styles usually 5, free, 2-2,5 mm 
long; stigmas capitate. Capsule ovoid, 5-6 mm 
long, 3 mm wide. 



A plant of damp places occurring in the eastern 
Transvaal, Swaziland, Natal Midlands and the eastern Cape. 

Transvaal. — 2430 (Pilgrims Rest): Mac Mac, Mudd 

Swaziland. — 2631 (Mbabane): Poliniane River, 
Compton 26489 (NBG). 

Natal. — 2831 (Nkandla): Nkandla Bush, Van der 
Merwe 2453. 2929 (Underberg): Ntabamhlope Research 
Station, West 634. 2930 (Pietermaritzburg): Balgowan, 
Mogg 3809; Howick Falls, Rogers 5212; Mooi River, 
Wood 4034; 6251 (K). 

Cape. — 3029 (Kokstad): 6 km W. of Bizana, Acocks 
12228. 3128 (Umtata): Baziya, Baur 582 p.p. (K). 3225 
(Somerset East): Bolus 319 (K). 3228 (Butterworth): Ken- 
tani, Pegler 117. 

Apparently Keller overlooked H. natalense Wood & 
Evans (1897) and described H. woodii (1923) basing it on 
the same Wood gathering. The latter is therefore a super- 
fluous name. 

6. Hypericum wilmsii Keller in Bull. 
Herb. Boiss. Ser. 2,8: 179 (1908) in clav.; 
Bredell in Bothalia 3: 579 (1939); Robson in 
Kew Bull. 12: 440 (1957); in F.Z. 1: 383 
(1961). Type: Transvaal, near Lydenburg, 
Wilms 136 (W, holo.; PRE, fragment!). 

Perennial herb, 6-20 cm high. Stems sev- 
eral to many, arising from a persistent taproot, 
procumbent, branched, terete, or somewhat flat- 
tened or 2-lined above, dark gland-dotted or 
eglandular. Leaves shortly petiolate, rarely al- 
most sessile; blade elliptic, ovate or obovate, 
6-10 mm long, 2-6 mm wide, apex rounded, 
base cuneate or rounded, nerves 3- or 4-paired, 

tertiary venation inconspicuous, with translu- 
cent^yellow gland dots and submarginal dark 
dots' petioles 1-2 mm long. Pedicels 3-10 mm 
long. Sepals unequal, oblong, 4,5-6 mm long, 
1-2 mm wide, apex obtuse to rounded, some- 
times apiculate, distinctly veined, yellow 
gland-dotted, and with 0-few dark dots along 
margin. Petals oblong, 5-8 mm long, 2-3 mm 
wide, dark gland-dotted along margin, yellow 
or orange. Stamens 18-30, in 3 or 4 groups, 
cohering at base; filaments c. 5 mm long; 
anthers 0,2-0, 3 mm long, each with dark gland 
at end of connective. Ovary ovoid, 3-4 mm 
long, 3- or 4-locular; styles 3 or 4, c. 2 mm 
long. Fruit capsular, 3- or 4-valved, erect. 

Occurring in moist areas and on mountain slopes in the 
Transvaal, Orange Free State, Lesotho and the Cape. Also 
in Rhodesia and Malagasy. 

Transvaal.— 2530 (Lydenburg): Lydenburg, Wilms 

O.F.S. — 2926 (Bloemfontein): Thaba ’Nchu, Roberts 
2350; 2913. 

Lesotho. — 2927 (Maseru): Ha-mvya-pela Mt., 
Likhoele, Mafeteng District, Dieterlen 1222; Catai River, 
Mafeteng District, Dieterlen 1293. 

Cape. — 3026 (Aliwal North): Doctor’s Drift, Gerstner 
137; Elandshoek, Bolus 153. 3126 (Queenstown): 

Queenstown, Galpin 1629. 3223 (Rietbron): Palmietfon- 
tein, Acocks 21770. 3225 (Somerset East): Cradock, 
Brynard 291. 

Differing from H. natalense in its shorter, procumbent 
stems and shortly petiolate leaves. H. nigropunctatum 
Norlindh, described from Rhodesia, is a synonym. 

5199 2. GARCINIA 

GarciniaL., Sp. PI. 1: 443 (1753); Gen. PI. ed. 5: 202 (1754); DC., Prodr. 1: 560 (1824); Benth. & 
Hook.f., Gen. PI. 1,1: 174 (1862); Harv., Gen. PI. ed. 2: 26 (1868); Oliv. in F.T.A. 1: 164(1868); 
Keller in Pflanzenfam. ed. 2, 21: 21 1 (1929); Phill. , Gen. ed. 2: 506 (1951); Robson in F.Z. 1: 396 
(1961). Type species: G. mangostana L. 

Trees or shrubs, usually with yellow juice, sometimes with leafy branches modified into 
thorns. Leaves opposite or in whorls of 3, sometimes large, usually entire, coriaceous, venation 
usually prominent, often with translucent glandular canals and brownish resin canals. Flowers 
terminal or axillary, solitary or in few- to many-flowered cymes, dioecious or polygamous, rarely 
bisexual. Sepals 4 (5), decussate. Petals 4 (5), longer than the sepals. Male flowers: Stamens 
indefinite, free or all connate in (3) 4 or 5 (-9) fascicles (bundles) with each fascicle forming a solid 
staminal column bearing free or partially fused filaments above, sometimes with a whorl of sterile 
stamen-fascicles (fasciclodes) alternating with the stamen-fascicles or forming a cushion in which 
the stamens are inserted; anthers opening by a circular rim or longitudinal slits. Female or bisexual 
flowers: usually with 4 (5) stamen- or staminode-fascicles, similar to those of the male flowers, but 
smaller and with fewer members, sometimes with fasciclodes, free or fused together in a ring at the 



base of the ovary. Ovary 2-many-locular with 1 ovule per loculus; styles absent; stigma sessile, 
peltate, lobed, smooth or tuberculate. Fruit baccate. Seeds 1-4, large, arillate; embryo with minute 

A large genus of about 400 species chiefly tropical, especially Asia, and South Africa. The genus was named in honour 
of Lawrence Garcin, an Englishman, who travelled and collected in India in the 18th century. 

Some species of the genus yield commercial products, e.g. G. mangostana L. — mangosteen fruits and G. morella 
Desr. — gamboge, but none of these is apparently cultivated in Southern Africa. 

Flowers terminal at ends of branches; staminal or staminodial fascicles present; leaves opposite; petioles 8-18 mm 

long 1 . G. gerrardii 

Flowers in fascicles in axils of older leaves on old wood; staminal or staminodial fascicles absent; leaves 3-whorled; 

petioles 4-8 mm long 2. G. livingstonei 

1. Garcinia gerrardii Harv. ex Sim, For. 
FI. Cape Col. 141, 1. 13 (14 May 1907); Henkel, 
Woody PI. Natal 126 (1934); Robson in F.Z. 1; 
398 (1961); Codd in Bothalia 8: 174 (1964); 
Bamps in Bull. Jard. Bot. Nat. Belg. 40: 282 
(1970). Type: Natal, Umvoti District, Gerrard 
1181 (BM!; K!). 

G. gerrardii Harv., Gen. S. Afr. PI. ed. 2: 26 (1868), 
nom. illegit. G. natalensis Schltr. in Bot. Jahrb. 40: 93 (21 
May 1907). Type: Natal, "in bewaldeten Schluchten am 
Umtwalumi bei Fairfield", Rudatis 150 (not traced). G. 
transvaalensis Burtt Davy in Kew Bull. 1924: 229 (1924); 
FI. Transv. 1; 252 (1926). Type: Transvaal, Rimers Creek, 
Barberton, Galpin 1202 (BOL!; K!; NH!; PRE!). 

Shrub or tree up to 13 m high; stems 
acutely angled or winged by decurrent petioles, 
with yellow or reddish brown latex; bark light 
grey. Leaves opposite; blade elliptic to broadly 
elliptic, 7-13 cm long, 3-8 cm wide, apex 
acute, mucronate with a recurved point, base 
broadly cuneate to rounded, midrib projecting 
below, lateral veins 9-13, Vi-angled, irregular, 
curved, looping near margin, margin subrevo- 
lute and callose, coriaceous, glossy on both 
surfaces, dark green above, paler below; petiole 
0,8- 1,8 cm long, furrowed, clasping. Flowers 

3- 9, terminal at ends of branchlets, subtended 
at base by small ovate-triangular bracts. Male 
flowers : Sepals 4, unequal, opposite and decus- 
sate, ovate-triangular, 1,5-2 mm long, 2 mm 
wide, keeled, mucronate (with brown tip), pale 
green with brown mottling (teste Scheepers 
1238). Petals broadly ovate to subrotund, 6-8 
mm long, 6-7 mm wide, concave, white. Stam- 
inal fascicles 4, 5,5 mm long, 1,5 mm wide, 
alternating with united rugose globose fasci- 
clodes; stamens 10-16; filaments free or partially 
united, 0,5-1 mm long; anthers 0,25 mm long, 
0,5 mm wide. Female flowers : Staminodial 
fascicles c. 1,5 mm long, 0,3 mm wide, alternat- 
ing with rugose globose fasciclodes; filaments 
united almost to apex. Ovary globose, 3- or 

4- locular, 2,6 mm long, 2,4 mm wide, sur- 

mounted by fleshy 3- or 4-lobed stigma. Berry 
subglobose, 1,8-3,25 cm diam., turning orange 
when ripe; seeds 1 or 2, subglobose, 1,4— 1,5 cm 

A constituent of forest, sometimes on streambanks, in 
the eastern Transvaal, Swaziland, Natal and the eastern 
Cape. Said to be particularly common in the Egossa Forest 
in Pondoland and in the Ngoye Forest in Zululand (Palmer 
& Pitman, Trees S. Afr., 1973). 

Transvaal. — 2531 (Komatipoort): 26 km from Barber- 
ton, Scheepers 1238; Rimers Creek, Galpin 1202. 

Swaziland. — 2531 (Komatipoort): Devils Bridge, 
Dlamini s.n.; Compton 31496. 

Natal. — 2830 (Dundee): Qudeni Forest, Edwards 2660. 
2831 (Nkandla): Eshowe, Gerstner 2541; Mtunzini Dis- 
trict, Wells & Edwards 29. 2930 (Pietermaritzburg): Kloof 
Nature Reserve, Moll 3256; Inanda, Wood 1082 (BM; K). 
2931 (Stanger): Hawaan Forest, Moll 3199. 3029 

(Kokstad): Bizana, Miller 4420. 3030 (Port Shepstone): 
Tributary of Uvongo River, Cooper 25. 3129 (Port St. 
Johns): Umsikaba, Strey 1012. 3130 (Port Edward): edge of 
Umtamvuna River Gorge forest. Cooper 103. 

Known as the Forest Mangosteen, um Bindi (X) and isi 
Binda (Z). Sim (l.c., 141) states that the saplings are used as 
whipsticks. The fruits are edible. 

2. Garcinia livingstonei T. Anders, in J. 
Linn. Soc. (Bot.) 9: 263 (1866); Oliv. in F.T.A. 
1: 165 (1868); Gibbs in J. Linn. Soc. (Bot.) 37: 
430 (1906); Sim, For. FI. P. E. Afr. 15, t.4 
(1909); Bak.f. in J. Linn. Soc. (Bot.) 40: 26 
(1911); Eyles in Trans. R. Soc. S. Afr. 5: 421 
(1916); Henkel, Woody PI. Natal 165 (1934); 
Brenan, Checklist Tang. Terr. 241 (1949); Keay 
in F.W.T.A. ed. 2,1: 294 (1954); Robson in 
F.Z. 1: 400 (1961). Type: specim. cult. hort. 
bot. Calcutta ex Mozambique, “In rupibus 
schistosis prope flumen Zambesi”, Kirk (CAL, 

G. angolensis Vesque in A. DC., Monogr. Phan. 8: 335 
(1893); R.E. Fr., Schwed. Rhod.-Kongo-Exped. 1: 151 
(1914); Burtt Davy, FI. Transv. 1: 252 (1926); Staner in 
Bull. Jard. Bot. Brux. 13: 120 (1934); Exell & Mendonpa in 
C.F.A. 1,1; 127 (1937). Type: Angola, Welwitsch 1047/48 
(K, iso!). G. baikieana Vesque, l.c. 336 (1893). Type: 
Nigeria, Barter 848 (K! ; P). G. ferrandii Chiov. in Stefa- 
nini & Paoli, Result. Sci. Miss. Stefanini-Paoli Somal. Ital. 
1: 26 (1916). Syntypes: Somalia, Paoli 847; 1012 (FI?). 



Fig. 5. — 1, Garcinia livingstonei, branch with bisexual flowers, x 2 h (Harrison 144); a, branch with male flowers, x 2 h 
(Stephen, Van Graan & Schwabe 1020); b, bisexual flower, x 10 (Harrison 144); c, male flower, x 10 (Stephen, 
Van Graan & Schwabe 1020); d, fruit, x 2 h (Stephen 27). 



Shrub or tree up to 12 m high, narrowly 
triangular when young, becoming bushy and 
virgate; branchlets 3-whorled, thick, flattened 
or angled towards apex; bark moderately 
smooth, grey with shallow furrows producing a 
chequered effect (teste De Winter & Vah- 
rmeijer 8572); sap yellow, thick. Leaves 
3-whorled, rarely 4- or opposite; blade elliptic, 
ovate or obovate to broadly so, 6-11 cm long, 
3-5,5 cm wide, apex acute to rounded, mucro- 
nate, base broadly cuneate, rounded or cordate, 
midrib projecting more on lower side, lateral 
veins 10-12 (-20), terminating in margin, ter- 
tiary venation variously conspicuous, margin 
entire or crenate, thickened, coriaceous, young 
leaves bright red; petiole 4—8 mm long, chan- 
nelled above, transversely rugose. Flowers 
polygamous, in fascicles of 5-15 or more, in 
axils of older leaves on old wood; pedicels 
0,8- 1,3 cm long, varying in thickness. Sepals 
4, unequal, in 2 opposite and decussate pairs, 
broadly elliptic to orbicular or triangular to 
transversely ovate, 1-3,5 mm long, 2-3 mm 
wide, concave. Petals 3-7 (9), ovate, obovate 
or orbicular, 4,5-5 mm long, 3-4 mm wide, 
concave, greenish white to cream or pale yellow 
with orange or reddish longitudinal glandular 
lines. Male flowers with numerous free stamens 
inserted on a fleshy cushion formed by the 
united fasciclodes. Bisexual (and female?) 
flowers with fewer stamens inserted in a fleshy 
fasciclodal ring below the ovary; ovary ovoid to 
globose, 2(3)-locular, 1,5 mm long, 2 mm wide. 

surmounted by fleshy bilobed stigma. Berry 
subglobose, 1-4 cm diam., yellowish red or 
orange, l-2(3)-seeded. Seeds piano-ovoid, 1-2 
cm long. Fig. 5. 

Found in scrub, open woodland and forest, often 
riverine, in the Transvaal, Natal, Swaziland, Botswana, 
Okavango, Caprivi Strip and tropical Africa. 

S.W.A. — 1723 (Singalamwe): E. of Kwando River, Cur- 
son 936. 1724 (Katima Mulilo): Ngoma, Killick & Leistner 
3322. 1821 (Andara): Andara Mission, De Winter 4238. 

Transvaal. — 2229 (Waterpoort): “Trekpad”, Vivo, 
Vahrmeijer 1786. 2231 (Pafuri): Pafuri Camp, Code! & 
Dyer 4625. 2329 (Pietersburg): Louis Trichardt, Stayt 4. 
2531 (Komatipoort): Malelane, Codd 4362; Van der Schijff 

Swaziland. — 2631 (Mbabane): Usutu Pont, Hlatikulu, 
Dlamini s.n. 2731 (Louwsburg): Hluti, Pole Evans 3392(1). 

Natal. — 2632 (Bela Vista): Maputa Region, Vahrmeijer 
1204; Maputa to Kosi Nature Reserve, Edwards 2591 . 2731 
(Louwsburg): banks of Black Umfolozi River, Nongoma, 
Codd 1936. 2732 (Ubombo): 13 km N.E. of Mseleni 
Mission Station, De Winter & Vahrmeijer 8572. 2832 
(Mtubatuba): Hluhluwe Game Reserve, Ward 2948; Palm 
Ridge Farm, Harrison 144. 

An extremely variable species, but readily distinguish- 
able from G. gerrardii (see key). According to a note on De 
Winter 4238, G. livingstonei is easily distinguished from 
other trees in the Okavango by its narrowly triangular shape 
and blue-green appearance. Palmer & Pitman (1973) state 
that “the general effect when the tree is bare of leaves is of 
a light-creamy grey or dark grey shape, all angles and 
points”. The flowers are said to be sweetly scented and the 
fruits, which have a pleasant acid taste, are relished by 
humans and animals. The wood is light-coloured and 
semi-hard and is used for fencing posts and boxes and 
sometimes furniture. The common names are Livingstone 
Garcinia, African Mangosteen and uPhimbi (Z). 


by A. A. Obermeyer 

Annual herbs or low shrublets with the branches either deciduous from a persistent woody 
crown, or perennial, often aromatic. Leaves decussate (rarely verticillate), simple, sessile or 
subsessile, serrate or rarely entire; stipules paired, interpetiolar, small, deciduous or persistent. 
Flowers axillary, solitary or in dichasia, pedicellate or subsessile, bisexual, actinomorphic, 
persistent; bracts resembling reduced stipules or 0. Sepals 2-5, free or fused at the base, imbricate 
with the outermost somewhat longer than the others. Petals 3-5, free, imbricate, shorter or longer 
than the sepals. Stamens diplostemonous or the inner whorl absent; filaments of outer whorl often 
expanded below, those of inner whorl shorter, not, or somewhat expanded, anthers bilocular, 
opening by longitudinal slits, introrse, versatile, dorsifixed. Gynoecium syncarpous, superior, 
3-5-locular, placenta axile, styles 3-5, free with apical, usually swollen stigmas, ovules numerous, 
attached to a convex placenta with the axis protruding and subulate above. Capsule septifragal; 



seeds subcylindrical, obtuse, straight or somewhat reniform to horseshoe-shaped, smooth or 
reticulate, exendospermous. 

Genera 2 with about 30 species. Cosmopolitan, inhabiting damp areas such as marshes, riverbanks or temporary pools; 
a few Bergia species are found in grassland communities. 

Flowers 5-merous; sepals with a prominent, firm midrib and membranous margin; ovary 

ovoid or globose, attenuate into short or long styles; herbs or shrublets 1. Bergia 

Flowers 3 (rarely 2)-merous; sepals without a prominent midrib, texture homogeneous; 
ovary depressed-globose with minute styles; small, creeping, soft herbs, usually 
hygrophytic 2. Elatine 

5230 1. BERGIA 

Bergia L., Mant. PI. Alt. 152, No. 1309 (1771); Harv. inF.C. 1: 1 15 (1860); Benth. & Hook., Gen. 
PI. 1; 162 (1862); Niedenzu in Pflanzenfam. 3, 6: 281 (1895); ed. 2, 21: 273 (1925); Phill., Gen. ed. 
2: 507 (1951); Hutch., Fam. Flow. PI. ed. 2, 1: 428 (1959). Type species: B. capensis L. 

Erect shrublets or with annual repent, decumbent or ascending branches from a woody crown, 
or annuals, glabrous, pilose or glandular-pubescent, often aromatic. Roots of the hygrophilous 
plants usually with a spongy outer covering. Stems and branches woody or herbaceous. Leaves 
decussate or rarely verticillate; lamina linear to ovate or obovate, margin entire or serrate, the teeth 
usually gland-tipped, pointing upwards. Flowers actinomorphous, small. Sepals 3-5 (5 in South 
African species), midrib prominent, margin membranous. Petals 3-5 (5 in South African species), 
delicate, white, pink or mauve. Stamens 10 (in South African species, except for B. ammannioides 
which has 5). Gynoecium 5-locular (in South African species) bearing 5 short or long, patent styles 
with apical, usually capitate, stigmas (in older flowers these curve outwards and touch the stamens; 
probably self-pollinated), ovules many. Capsule with the 5 valves breaking away from the hard 
thin dissepiments, the placentas bulging or recurved to horn-shaped, covered with the short 
hardened funicles; above the axis is prolonged into a subulate tip. Seed very small, subcylindric to 
somewhat reniform, usually obtuse above and below, tesselated or smooth, yellow to brown or 

A cosmopolitan genus with about 25 species; 10 of these occur in Southern Africa. Those species which are 
hygrophilous may well be distributed by birds or floodwaters for the seeds are very small. 

The genus was named in honour of Petrus Jonas Bergius a professor in Stockholm and author of Descriptiones 

plantarum ex Capite Bonae Spei, 1767, an early flora on Cape plants. 

An anatomical study of this genus might prove interesting. 


Flowers 1-2 in the leaf axils, on long pedicels (up to 15 mm): 

Petals longer than the sepals; flowers distributed all over the branches 1. B. anagalloides 

Petals shorter than the sepals; flowers aggregated towards the apices of the branches 2. B. polyantha 

Flowers several to many in the leaf axils; pedicels short (up to 7 mm): 

Stems erect, rarely branched, from a creeping rhizome, succulent, glabrous; valves of capsule flat and recurved at 

dehiscence; usually an aquatic 3. B. capensis 

Stems prostrate or ascending, numerous, radiating from a central base; valves of the capsule usually hollow at 
dehiscence; wet, sandy places: 

Stamens 10; plants glandular-pubescent; petals longer than the sepals: 

Sepals spathulate, midrib and margins with some long filiform processes; styles as long as the 

ovary 4. B. spathulata 

Sepals linear, unequal in length, resembling the stipules and bracts, fimbriate; style sessile or nearly 

so 5. B. glutinosa 

Stamens 5; plants pilose with whitish hairs rarely mixed with some glandular pubescence; petals shorter than 

the sepals 6. B. ammannioides 




Leaves small, obovate (c. 5 X 3 mm), soft; plants forming dense, woody mats; intemodes short, often almost 

invisible; flowers solitary; a Cape endemic 7. B. glomerata 

Leaves larger, linear to ovate, firm; plants with spreading or erect, usually annual branches from a persistent woody 
crown or branches perennial; intemodes well developed; flowers several to many in each axil; summer rainfall 

Stems repent, pilose; leaves about 1 cm long, narrowly ovate, coriaceous; flowers c. 2,5 mm long, sepals short, 

spine-tipped; styles short 8. B. pentherana 

Stems erect or ascending, sparsely strigose or scabrid; leaves coriaceous or soft, 1,5-3 cm long; flowers c. 

3-6 mm long, sepals acuminate; styles long: 

Leaves linear, usually glabrous, hard and shiny when dry with a swollen margin and prominent veins, the 6-9 

teeth widely spaced 9. B. decumbens 

Leaves narrowly ovate, glandular-pubescent, soft, margin and nerves not prominent, the many teeth close 

together, usually double-serrate; stems annual or if perennial, forming untidy bushes 10. B. salaria 

1. Bergia anagalloides E. Mey. ex Fenzl 
in Ann. Naturh. Mus. Wien 1: 344 (1836); 
Walp., Repert. 2: 786 (1843); Harv. in F.C. 1: 
116 (1860); Thes. Cap. 2: 22 t. 133 (1863); 
Niedenzu in Pflanzenfam. 3, 6: 281 (1895), ed. 
2, 21: 274 (1925); Roessler in F.S.W.A. 92: 3 
(1968). Type: Cape, on the bank of the Gariep 
(Orange River) below 200 feet, Drege 2957 
(W, holo.!; PRE, photo.). 

B. alsinoides Holzhammer in Mitt. Bot. Staatssamml. 
Miinchen 1: 335 (1953). Type: S.W. Africa, 5 km E. of 
Weissenfels, Dinter 8054 (M, holo.; PRE). 

Small, prostrate annual with many radiat- 
ing branches, somewhat woody at the base. 
Roots spongy, with a long taproot (fide Bryant). 
Stems slender, smooth or with indistinct scabrid 
ridges. Leaves narrowly obovate, subpetiolate, 
up to 3 cm long and 6 mm broad, often smaller, 
serrulate, nerves distinct below, soft in texture; 
stipules minute. Flowers solitary, axillary, 
evenly distributed along the stems, on firm 
pedicels up to 15 mm long; bracts minute or O. 
Sepals narrowly ovate, acute, 2,5 mm long, 
hollow, green with the membranous margins 
often pink. Petals longer than sepals, 4 mm, 
oblong, obtuse, bright pink. Ovary ovoid with 
short arcuate styles. Capsule ovoid, 3 mm long; 
seed oblong, 0,4 mm, tesselate, yellow to gol- 
den brown. Fig. 6: 2. 

Recorded from South West Africa, the north-western 
Cape and south-western Orange Free State, in damp places, 
usually calciphilous. 

S.W. A. — 2316 (Nauchas): 5 km E. of Weissenfels, 
Dinter 8054. 

Cape. — 2624 (Vryburg): Armoedsvlakte, Mogg 7946. 
2723 (Kuruman): Bathlaros near Kuruman, Silk 64. 2818 
(Warmbad): Ramans Drift, Orange River, Schlechter 
11453. 2824 (Kimberley): Holpan, Acocks 2463. 2920 
(Boom Rivier): Seweperd, S.E. of Pofadder, A cocks 15076. 
2922 (Prieska): on sandy banks of Orange River, Bryant 

783. 2923 (Douglas): farm Mazelsfontein, junction of 
Orange and Vaal Rivers, Anderson 610. 3024 (De Aar): 
near Potfontein, Acocks 2413. 

O.F.S. — 2925 (Jagersfontein): Fauresmith, Waagkop, 
Henrici 3910. 

2. Bergia polyantha Sond. in Linnaea 23: 
16 (1850); Harv. in F.C. 1: 116 (1860); Oliv. in 
F.T.A. 1: 153 ( 1868); Niedenzu in Pflanzenfam. 
3, 6: 281 (1895); ed. 2, 21: 273 (1925); Burtt 
Davy in FI. Transv. 1: 147 (1926); Exell & 
Mendonca in C.F.A. 1, 1: 118 (1937); Wild in 
F.Z. 1: 375 (1961); Roessler in F.S.W.A. 92: 4 
(1968). Type: Orange Free State, Rhinosterkopf 
by the Vaal and Sand Rivers, Burke & Zeyher 
540 (K, holo.). 

B. integrifolia Dinter ex Holzhammer in Mitt. Bot. 
Staatssamml. Miinchen 1: 335 (1953); Roessler in F.S.W.A. 
92: 4 (1968). Type: S.W. Africa, Grootfontein, Grosshuis, in 
a swamp, Dinter 7381 (M, holo.; K; PRE). 

Glabrous annuals with many radiating de- 
cumbent branches up to 20 cm long. Roots 
numerous, swollen. Stems terete, smooth. 
Leaves narrowly obovate, up to 3 cm long and 1 
cm broad, margin entire or faintly serrulate, 
soft; stipules small. Flowers 1-2 in the leaf 
axils, on thin pedicels up to 2 cm long, aggre- 
gated apically where the intemodes and 
pedicels are shorter. Sepals narrowly ovate- 
acuminate, 2 mm long, green with white to pink 
margins. Petals shorter than sepals, obovate, 
1,5 mm long, pink or mauve. Stamens with thin 
filaments. Ovary ovoid, styles very short, 
arched over the ovary. Capsule globose, 
placenta often sickle-shaped, echinate; seeds 
oblong-cylindrical, 0,5 mm, tesselate, brown, 
shiny. Fig. 6: 3. 

Recorded from South West Africa, the western Orange 
Free State, northern Cape, Botswana, Rhodesia, Mozam- 
bique, Zambia and Angola; in moist places. 


Fig. 6. — 1, Bergia glomerata, x 1; la, leaf with apical gland on lower surface, X 4; lb, flower with carpels, X 8; lc, sepal, 
X 10; Id, petal, x 10; le, stamen, x 10; If, gynoecium, x 10; lg, seed, x 22. 2, B. anagalloides, leaf, x 1; 2a, 
stipule, X 14. 3, B. polyantha, leaf, X 1; 3a, stipule, x 8. 4, B. capensis, leaf, x 1; 4a, stipule, x 11. 5, B. 
spathulata, leaf, x 1; 5a, stipule, X 5. 6, B. glutinosa, leaf, x 1; 6a, stipule, x 5. 7, B. ammannioides, leaf, x 1; 7a, 
stipule, x 5. 8, B. glomerata, leaf, x 1; 8a, stipule, x 5. 9, B. pentherana, leaf, x 1; 9a, stipule, x 3. 10, B. 
decumbens, leaf, x 1; 10a, stipule, x 4. 11, B. salaria, leaf, x 1; 11a, stipule, x 5 (Semi-diagrammatic). 



S.W.A. — 1712 (Posto Velho): on the Kunene Riverbank 
at Otjinungua, Giess 8871. 1713 (Swartbooisdrift): Kunene, 
riverbank at Epupa Falls, Giess & 1 Viss 3233; Otjitambi, in 
the Hondoto River on the road to Swartbooisdrift, Giess 
3126. 1813 (Ohopoho): Kaoko Otavi, De Winter & Leistner 
5557. 1821 (Andara): Gam, Story 5115. 1918 (Grootfon- 
tein): Grosshuis, Dinter 7381 . 2016 (Otjiwarongo): Omat- 
jenne Experimental Farm, Pfeiffer s.n. 2115 (Karibib): 
Klein Ameib, Dinter 7055; 7056. 2117 (Otjosondu): Ot- 
josondu Farm, KAR 36, Rote Berge, Giess 3436. 

3. Bergia capensisL., Mant. 241 (1771); 
Milne-Redh. in Kew Bull. 1948: 450 (1948); 
Subramanyam, Aquatic Angiosperms, India, 
Bot. Mon. No. 3: 10, f.6 (1962); Cook in FI. 
Europ. 2: 295 (1968); Roessler in F.S.W.A. 92: 
2 (1968); Verdcourt in F.T.E.A. Elatinaceae: 3, 
f. 1, 1-7 (1968). Type: presumably from Asia, 
not from the Cape (LINN 597. 1 holo. ; PRE, 

B. aquatica Roxb., PI. Corom. 2: 22 ( 1798), nom. illegit. 
B. verticillata Willd., Sp. PI. 2: 770 (1799); F.T.A. 1: 152 
(1868), nom. illegit. 

B. sessiliflora Griseb., Cat. PI. Cub. 40 (1866); 
Monachino in Phytologia 5: 184 (1955). Type from Cuba. 

Elatine luxurians Del., Descr. Egypt. Hist. Nat. 2: 216 
(1813), nom. illegit. 

Succulent, glabrous aquatic annuals with 
erect stems from a creeping rhizome. Roots 
numerous at each rhizome node, stout, with 
many long thin side roots. Stems up to 25 cm 
long, and up to 7 mm in diam., unbranched or 
with a few short side branches, succulent. 
Leaves reduced below, upper fairly close to- 
gether, narrowly ovate, up to 4 cm long and 1 
cm broad, attenuate below, shallowly serrate, 
thin; stipules small. Flowers small, in many- 
flowered, tight clusters, sessile or very shortly 
pedicellate. Sepals narrowly ovate, 1,5 mm 
long, membranous with entire margins. Petals 
narrowly ovate, slightly longer, pale lilac. Sta- 
mens 10. Ovary globose, styles short. Capsule 
globose, the valves flattened and recurved at 
dehiscence; seeds subcylindrical, straight or 
slightly bent, 0,8 mm long, tesselate, shiny 
yellow brown. Fig. 6: 4. 

Recorded from South West Africa and the eastern 
Transvaal; also in tropical and subtropical Africa and Asia. 
An aquatic often found in semi-permanent rock pools. 

S.W.A. — 2115 (Karibib): Umduasbank, Dinter 7025. 
2417 (Mariental): farm Haribes, Leippert 4721 . 

Transvaal. — 2531 (Komatipoort), Kruger National 
Park, Shabin Kop near Pretorius Kop, Codd 6054; 
Malelane byroad, 9 km from Kemps cottage, Ihlenfeldt 
2386; Mbiamide, Van der Schijff 1967. 

4. Bergia spathulata Schinz in Bull. 
Herb. Boiss. 6: 526 (1898); Niedenzu in Pflan- 
zenfam. ed. 2, 21: 274 (1925); Exell & Men- 
donpa in C.F.A. 1: 119 (1937); Roessler in 
F.S.W.A. 92: 4 (1968). Type: S.W. Africa, 
Olukonda, Rautanen 221 (Z, holo.). 

B. erythroleuca Gilg in Warb., Kunene-Samb. Exped. 
308 (1903). Type: Angola, on the left bank of the Kunene 
River near Soba Gongo, Baum 1 10 (HBGt). 

Small prostrate or ascending, glandular- 
pubescent, aromatic annuals with numerous 
radiating branches, up to 40 cm tall. Stems thin, 
wiry, dark red. Leaves narrowly obovate, at- 
tenuate below, up to 2 cm long and 4 mm wide, 
minutely serrate, thin; stipules small, fimbriate. 
Flowers in dense rounded compound cymes 
present in every axil, the brown sepals and 
white petals giving the clusters a mottled ap- 
pearance; bracts resembling stipules, up to 4 
mm long; pedicels up to 2 mm. Sepals narrowly 
spathulate, 3 mm long, margin and midrib 
fimbriate, green, apex red or brown, apiculate. 
Petals narrowly obovate-acuminate, about 4 
mm long, white or violet. Stamens 10. Ovary 
ovoid, 5-furrowed, papillate, red, the styles 
about as long as the ovary, erect. Capsule 
ovoid, red, papillate; seed 0,75 mm long, tesse- 
late. Fig. 6: 5. 

Known only from Angola and northern South West 
Africa, in moist places after rains. 

S.W.A. — 1713 (Swartbooisdrif): 48 km N. of Ohopoho 
on the road to Ruacana, Giess & Leippert 7547. 1817 
(Tsintsabis): farm Tsintsabis, Dinter 7668. 

Rautanen informed Schinz that the local natives sowed 
the seeds of this plant mixed with those of their water- 
melons hoping that the sweet honey-like fragrance would 
enter their melons. 

5. Bergia glutinosa Dinter & Schulze- 
Menz in Notizbl. Bot. Gart. Berk 15: 453 
(1941); Roessler in F.S.W.A. 92: 3 (1968). 
Type: S. W. Africa, Grootfontein, Wackies, 
Schoenfelder sub herb. Dinter 7223a (B, holo.; 
M; PRE). 

Small annuals with numerous radiating 
branches, scabrid, glandular-pubescent. Stems 
thin, wiry. Leaves narrowly ovate, up to 4 cm 
long and 12 mm broad, shortly petiolate, ser- 
rate, thin; stipules linear, 1,5 mm long, mem- 
branous, fimbriate. Flowers in several-flowered 
cymes, shortly pedicellate. Sepals linear, nearly 
twice as long as the petals, resembling the 
bracts, unequal in length, the outer longest,' up 
to 4 mm long, green, fimbriate, margin mem- 
branous below. Petals linear, about 2 mm long, 



pink. Stamens 10. Ovary globose with the 
styles sessile or nearly so. Capsule globose; 
seeds 0,8 mm long, smooth or with indistinct 
markings. Fig. 6: 6. 

Known only from two gatherings in northern South 
West Africa; in moist places. 

S.W.A. — 1918 (Grootfontein): Wackies, Schoenfelder 
sub herb. Dinter 7223a; Naruchas (Nurugas, GR 923/924), 
Dinter 7223. 

Wild, in Flora Zambesiaca 1: 376 (1961), places a 
collection by Drummond & Seagrief 5186 from Botswana, 
Nata River, under this apparently annual species but I prefer 
to regard this plant as belonging to the perennial B. salaria 
Brem. The sessile flowers with the long, linear, fimbriate 
sepals and the globose capsule with short styles distinguish 
B. glutinosa from B. salaria, which has pedicellate flowers 
with ovate-acuminate subulate sepals only slightly longer 
than the petals and an ovoid capsule topped with long styles. 

A collection ( Mitchell 922) from Lake Kariba appears 
to be nearest B. glutinosa but is apparently a perennial and 
somewhat larger in all respects. This will need further 

6. Bergia ammannioides Hevne ex Roth, 
Nov. PI. Sp. 219 (1821); DC., Prodr. 1: 390 
(1824); Oliv. in F.T.A. 1: 152 (1868), as 
ammanoides', Niedenzu in Pflanzenfam. 3, 6: 
281 (1895); Engl., Pflanzenw. Afr. 3, 2: 524, 
t. 237, fig. K-R (1921); Niedenzu in Pflanzen- 
fam. ed. 2, 21: 274 (1925); Keay in F.W.T.A. 
ed. 2, 1: 128 (1954); Wild in F.Z. 1: 375, t.72, e 
(1961), pro parte; Roessler in F.S.W.A. 92: 3 
(1968); Verdcourt in F.T.E.A. Elatinaceae: 3, 
fig. 1, 8 (1968). Type: India, Heyne (K, holo.). 

Small prostrate annual with numerous thin 
radiating branches, minutely and sparsely 
pilose, especially in the axils, giving it a whitish 
appearance. Stems thin with long intemodes. 
Leaves narrowly ovate to obovate, attenuate 
into a short petiole, variable in size, 15-25 mm 
long and 3-5 mm wide, margin serrate, 
coriaceous, glabrous or sparsely pilose; stipules 
linear, acuminate, about 2 mm long, ciliate, 
membranous. Flowers small, c. 2 mm, in dense 
globose, reddish cymes; pedicels up to 3 mm, 
pilose. Sepals linear-acuminate, 2 mm long, 
membranous with a broad green or red, pilose 
midrib, margin smooth. Petals narrowly ovate, 
1,5 mm long, white or pink. Stamens 5 (in 
South African specimens seen), short, repre- 
senting the outer circle. Ovary globose, styles 
short. Capsule with the valves often blackish 
red at the apex; seeds 0,5-0,75 mm long, 
tesselate, yellow. Fig. 6: 7. 

Recorded from the warmer drier regions of Africa, Asia 
and Australia, where it is found in damp places. In Southern 
Africa recorded from South West Africa; also in Rhodesia. 

S.W.A. — 1918 (Grootfontein): Abenab-Gunib, Dinter 
7380; Grootfontein, Dinter 7627; Ameib near Grootfon- 
tein, Schoenfelder S635. 2016 (Otjiwarongo): Okateitei, 
OTJ 157, Dinter 7844. 

Note: The glabrous specimens cited by Wild in Flora 
Zambesiaca 1: 375 (1961) belong to B. capensis L. 

7. Bergia glomerata L.f, Suppl. 243 
(1782); Thunb., Prodr. 82 (1794); FI. Cap. ed. 
Schult. 395 (1823); DC., Prodr. 1: 390 (1824); 
Harv. in F.C. 1: 116 (1860); Niedenzu in 
Pflanzenfam. 3, 6: 282 (1895); ed. 2, 21: 274 
(1925); Adamson in FI. Cape Penins. 588 
(1950). Type: Cape, near Swartkops River, 
Uitenhage, Thunberg (S, holo.; PRE, part of 

Perennial, woody prostrate glabrous plants 
forming mats often up to 1 m in diam., usually 
turning brown when dry. Stems thick, woody, 
with numerous short and long side branches, the 
intemodes often telescoped together, densely 
leafy. Leaves small, obovate, 5 mm long, 3 mm 
broad with a few teeth towards the rounded 
apex or edentate, glabrous, smooth; on the 
lower side with a small elongate cushion- 
shaped gland apically; stipules large, ovate, 
pectinate. Flowers sessile, small, c. 3 mm, few 
to several in axillary glomerules. Sepals oblong 
to lanceolate-acute, about 2,5 mm, firm. Petals 
oblanceolate, 3 mm long, white, folded below 
with the upper half turned inwards. Stamens 
with the outer filaments somewhat wider and 
longer. Ovary globose, red, papillate, styles 
erect, terete. Capsule 5-partite, ovoid; seeds 
allantoid, 0,75 mm long, tesselate, black or 
brown. Fig. 6: 1. 

A Cape endemic found around the coast or somewhat 
further inland, from Vanrhynsdorp in the west to the Albany 
district in the east, in temporary pools or damp places. 

Cape. — 3118 (Vanrhynsdorp): farm Liebendal, 1 1 km N. 
of Vredendal, Hall 3918. 3119 (Calvinia): Lokenburg, 
Acoc/cs 17577; 17467. 3318 (Cape Town): 27 km from 
Malmesbury to Hopefield, Marsh 646; N. of Tygerberg, 
Compton 20061. 3319 (Worcester): Worcester, Van Breda 
1056. 3322 (Oudtshoom): Kruispad, Compton 21772. 3325 
(Port Elizabeth): at the mouths of the Koega and Swartkops 
Rivers, Zeyher 1956; near the Swartkops and Gamtoos 
Rivers, Uitenhage, Zeyher 1768; Swartkops River and 
Koegaskop, Zeyher 513; Korsten commonage. Long 1141. 
3326 (Grahamstown): “Strowan” farm near Grahamstown, 
Britten 6575. 3420 (Bredasdorp): De Hoop Nature Reserve, 
Van der Merwe 1980; Bontebok Park, Maguire 820. 3421 
(Riversdale): Riversdale coast, Marloth 50; 8064. 

Of interest are the vestiges of the pectinate stipules 
found adnate to the bases of the bracts and outer sepals. 
From this it can be deduced that the thin transparent or 
white, often ciliate margins of the sepals are of stipular 
origin and now fused to the sepal. The elongate swollen 
dorsal apical gland has been observed in this species only; 
its function is unknown. 



8. Bergia pentherana Keissl. in Ann. 
Naturh. Hofm. Wien 15: 58 (1900)*; Niedenzu 
in Pflanzenfam. ed. 2, 21: 274 (1925). Type: 
Rhodesia, Wankie Game Reserve, Bulungweti 
pans (“Land der 1000 Vleys, Bolongeti or 
Bellumbeti”), Penther 2317 (W, holo. !; PRE, 

Bergia prostrata Schinz in Mem. Herb. Boiss. 20: 22 ( 15 
Oct. 1900); Engl., Pflanzenw. Afr. 3, 2: 524(1921); Wild in 
F.Z. 1: 376 (1961); Roessler in F.S.W.A. 92: 4 (1968). 
Type: S.W. Africa, Orumbo on the Nossob River, Dinter 
1335 (Z, holo.). B. palliderosea Gilg in Warb., Kunene- 
Samb. Exped. 308 (1903); Exell & Mendonpa, C.F.A. 1: 
1 18 (1937). Type: Angola, on the left bank of the Kunene 
River above Humbe, Baum 98 (HBG t). 

Prostrate perennial with long radiating an- 
nual branches, often rooting at the nodes, from 
a perennial crown; side branches many, short, 
prostrate or erect. Stems terete, pubescent. 
Leaves small, uniform in shape, but varying 
somewhat in size, narrowly ovate, 5-10 mm 
long and up to 3 mm broad, coriaceous, margin 
raised, laxly serrulate, nerves prominent below; 
stipules linear, serrulate. Flowers small, about 
2,5 mm long, in 4— 7-flowered axillary 
glomerules, pedicels about 2 mm, hairy. Sepals 
narrowly oblong, 2 mm long, acute, mucronate, 
firm. Petals ovate, 2,5 mm long, white or pink. 
Stamens 10, purple, with the outer filaments 
somewhat expanded and slightly longer than 
inner. Ovary ovoid, purple, styles fairly short 
with bulbous stigmas. Capsule 5-partite, 
globose with the valves hollow; seeds oblong- 
cylindrical, 0,75 mm tesselate, brown, usually 
1-few developed in each carpel. Fig. 6: 9. 

Recorded from Angola, northern South West Africa, 
Rhodesia, northern Cape, south western Transvaal and 
western Orange Free State, i.e. around the borders of the 
Kalahari; in sand, often in limestone areas; common around 
Kimberley and surrounding districts. 

S.W. A. — 1715 (Ondangua): 17 km E. of Oshikango, De 
Winter & Giess 7047. 1718 (Kuring-kuru): Omuramba 
Mpungu, Tsintsabis-Kuring-kuru road, De Winter 3875. 
1719 (Runtu); Tamsoe, S.E. of Runtu, Maguire 1633. 1917 
(Tsumeb): Otavi, Dinter 5235. 1918 (Grootfontein): 

Okavango, 16 km W. of Xeidang, Giess 10051. 2117 
(Otjasondu): Quickborn, Bradfield 449. 2218 (Gobabis): 
farm Hester, Merxmuller & Giess 1216; Okasewa, Dinter 

Transvaal. — 2526 (Zeerust): Panfontein Game Re- 
serve, Louw 2044. 2725 (Bloemhof): 20 km S. of 
Bloemhof, Acocks 12583. 

O.F.S. — 2726 (Odendaalsrus): Odendaalsrus, Goossens 

Cape. — 2624 (Vryburg): Armoedsvlakte, Mogg 8111. 
2625 (Delareyville): Setlagodi, Burst Davy sub PRE 11032. 
2723 (Kuruman): Esperanza, near Kuruman, Esterhuysen 
2244. 2824 (Kimberley): Barkley West, Pienaarsfontein, 
Acocks 154. 

9. Bergia decumbens Planch, ex Harv., 
Thes. Cap. 1: 15, t.24 (1859); F.C. 1: 116 
(1860); Oliv. in F.T.A. 1: 153 (1868); Niedenzu 
in Pflanzenfam. 3, 6: 281 (1896); ed. 2, 21: 274 
(1925); Engl., Pflanzenw. Afr. 3, 2: 524(1921); 
Marloth, FI. S. Afr. 2; 184, f. 123 (1925); Burn 
Davy, FI. Transv. 1: 147 (1926); Wild in F.Z. 1: 
377 (I960). Type: Transvaal, Magaliesberg, 
Burke & Zeyher s.n. (K, holo.; PRE; SAM). 

Small undershrubs 20-50 cm tall with 
many annual (or rarely perennial) erect and 
procumbent branches arising from a perennial 
woody crown, red when young, soon becoming 
lignified, glabrous or hispidulous; side branches 
short or repressed to form leafy fascicles. 
Leaves linear to broadly linear, 1,5-2, 5 cm 
long, 2-4 mm broad, uniform in shape, margin 
appearing swollen in the dried state with about 
4-9 (-12) widely spaced teeth, the veins fairly 
broad, consisting of elongated cells contrasting 
sharply with the rest of the lamina where the 
cells appear small and squared to rounded, 
leathery in texture, shiny; stipules small, linear, 
serrulate. Flowers small, c. 4 mm long, in 
2-8-flowered dichasia, pedicels pilose, c. 2 mm 
long. Sepals ovate-acuminate, 4—5 mm long, 
mucronate, margins laxly fimbriate. Petals 
slightly longer than sepals, narrowly obovate, 
white or pink. Stamens 10, filaments narrowly 
ovate-acuminate. Ovary ovoid-attenuate with 5 
long, clavate, arcuate styles. Capsule narrowly 
ovoid, the valves usually flattened and spread- 
ing at dehiscence; seeds 0,5 mm, black, tesse- 
late. Fig. 6: 10. 

Widespread and fairly common in grassland in drier 
parts of the Transvaal, eastern Botswana, Rhodesia and 

Transvaal. — 2329 (Pietersburg): near Pietersburg, 
Bolus 10900. 2330 (Tzaneen): Tzaneen, Rogers 12545. 
2428 (Nylstroom): Schaaphok, c. 13 km N. of P.O. Riet- 
bokspruit. Story 1637; Codd 2341; above Warmbaths, 
Smuts & Gillett 3321; Bolus s.n. 2429 (Zebediela): 
Pyramid Estate near Potgietersrus, Galpin 8990; Lunsklip, 
Maguire 1396. 2526 (Zeerust): Zeerust, Leendertz sub TRV 
11295. 2527 (Rustenburg): Rustenburg, Leendertz sub TRV 
9743. 2528 (Pretoria): Rietvlei Veld Reserve, Acocks 
11281. 2530 (Lydenburg): Crocodile River, Schlechter 
3912. 2626 (Klerksdorp): Lichtenburg, White’s quarry. 

* The exact date of publication could not be established. Since however the species appeared in the first of four parts of 
Ann. Naturh. Hofm. Wien 15 (1900), it may be assumed that it was published before October, the month in which Schinz’s 
Bergia prostrata appeared. 



Morris & Engelbrecht 1030. 2627 (Potchefstroom): Bos- 
kop, near Potchefstroom, Louw 1626. 2628 (Johannesburg): 
32 km S. of Heidelberg, Acocks 21024. 

The species possibly hybridises with B. salaria, which 
would account for the glandular pubescence found on plants 
collected in the northern Transvaal, northern Natal and 
southern Mozambique, e.g. Galpin M68, M289 from Mos- 
dene near Naboomspruit in the Transvaal, Strey & Moll 
3789 from near Maputa in northern Natal and Drummond 
7724 from Mozambique, etc. These specimens appear to be 
intermediate between these two taxa. This will need further 

10. Bergia salaria Brem. in Ann. Transv. 
Mus. 15: 248 (1933). Type: Transvaal, Sout- 
pansberg, Soutpan, Bremekamp & Schweick- 
erdt 227 (PRE, holo.). 

B. mossambicensis Wild in Bol. Soc. Brot. Ser. 2, 31: 93 
( 1957); in F.Z. 1: 375, t.72 f.C ( 1961). Type: Mozambique, 
Sul do Save, Guija district, Limpopo River, Gazaland 
Expedition sub TRV 15816 (SRGH, holo.; PRE). 

Small glandular-scabrid undershrubs with 
erect or decumbent annual stems from a woody 
crown; if perennial, the stems elongate, produce 
abbreviated side branches and adopt a more 
straggling habit. Leaves on the primary 
branches ovate, 2-4 cm long, 1-2 cm broad, 
acute, teeth many, close together, often 
double-serrulate, fairly soft in texture, veins 
narrow, with elongated cells, deciduous; on 
perennial stems the later leaves produced on the 
abbreviated secondary branchlets are smaller; 
stipules linear-acuminate, about 5 mm long. 
Flowers in 2-7-flowered cymes; pedicels 1-7 
mm long. Sepals ovate, subulate, 5-6 mm long, 

the broad midrib green, glandular-pubescent, 
the membranous margins red and white, den- 
ticulate. Petals shorter than sepals, broadly 
ovate-acute, 4-5 mm long, white, pink or pale 
mauve. Stamens 10 with the outer filaments 
somewhat longer and expanded below. Ovary 
5-lobed, ovoid, attenuate into the styles which 
are about as long as the ovary. Capsule ovoid, 
the valves hollow at dehiscence, thin; seeds 
oblong-cylindrical, 0,5 mm, tesselate, dark 
brown, shiny. Fig. 6: 11. 

Found in warmer parts of the Transvaal, Rhodesia and 
Mozambique, on sandy riverbanks or beside pans, possibly 
distributed by floods. 

Transvaal. — 2229 (Waterpoort): Dongola, farm Nekel 
620, Codd 4095. 2230 (Messina): Messina, Rogers 19303. 
2231 (Pafuri): Machai pan, Nwambiya, Brynard & Pienaar 
4264; Van der Schijff 5688. 2328 (Baltimore): Magalakwin, 
Smuts sub PRE 1980. 

As these plants, which inhabit the warmer parts of the 
Transvaal and neighbouring countries, often perennate, 
their appearance in time becomes more lax and straggly; the 
primary leaves are dropped and on the short side branches 
smaller leaves take their place. 

The type material of B. mossambicensis Wild, from 
the banks of the Limpopo River, appears to consist of young 
plants. It matches Codd 4095 from Dongola near Messina 
and links up with older specimens which match B. salaria 
from the Soutpan, not far from Dongola. B. salaria is also 
recorded from Rhodesia, e.g. Drummond 5762 from near 
Chiturupadzi store 40 km N.N.W. of the Bubye-Limpopo 
confluence, Phipps 2906 from the Nuanetsi district, 48 km 
S. of Chipanda pools, etc. Typical plants of B. decumbens 
and B. salaria can be distinguished easily but there are 
intermediates in areas where both species occur which seem 
to suggest that they hybridise (cf also note under B. 

5231 2. ELATINE 

Elatine L., Sp. PI. 1: 367 (1753); Gen. PI. ed. 5: 172 (1754); Benth. & Hook.f., Gen. PI. 1: 162 
(1862); Niedenzu in Pflanzenfam. 3, 6: 282 (1895); ed. 2, 21: 276 (1925); Hutch., Fam. Flow. PI. 
ed. 2, 1: 428, f. 27 1 (1959); Wild in F.Z. 1: 377 (1961). Type species: E. hydropiper L. 

Small herbaceous, glabrous, aquatic or semi-aquatic annuals. Leaves decussate or verticillate, 
margin entire or serrulate; stipules small, deciduous. Flowers solitary in the leaf-axils, sessile or 
nearly so, ebracteate, very small, often cleistogamous. Sepals 2-4, connate below, midrib 
indistinct, margin not membranous. Petals 2-4, free, larger than sepals, white or pink. Stamens as 
many as the petals or diplostemonous. Ovary globose, 3-4-locular, styles 3-4, minute, situated on 
the depressed apex of the ovary. Capsule globose, membranous, the dissepiments remaining 
attached to central axis as wings; seeds numerous, straight or bent to horseshoe-shaped, tesselate. 

Cosmopolitan with about 12 species occurring in tropical, subtropical and temperate regions as aquatics or in moist 
situations. One or possibly 2 species recorded from Southern Africa. 


Fig. 7. — 1, Elatine triandra, x 1; a, flower, x 50; b, seed, x 80 (Ward 2574). 



1. Elatine triandra Schkuhr, Bot. Handb. 
ed. 2, 1: 345, 1. 1096 (1808); Niedenzu in 
Pflanzenfam. 3, 6: 383 f. 131 K-L (1895); ed. 

2, 21: 276, 120 K-L (1925); Wild in F.Z. 1: 378 
(1961); Subramanyam, Aquatic Angiosperms 
India, Bot. Mon. No. 3: 10 (1962). Type from 

A small, soft creeping annual herb, rooting 
at the nodes. Leaves decussate, shortly petiol- 
ate, linear-oblong to narrowly ovate, up to 15 
mm long and 3 mm broad, apex emarginate, 
narrowed at the base into a short petiole; 
stipules membranous, deciduous. Flowers ses- 
sile or with pedicels up to 1 mm long. Sepals 3 
(rarely 2), narrowly ovate, 0,5 mm long. Petals 

3, broadly ovate, 1-1,5 mm long, white or pink. 
Stamens 3, shorter than petals. Ovary 3-locular, 

globose, styles very short. Capsule membran- 
ous; seed slightly curved, oblong-terete, 0,5 
mm, tesselate, light brown, shiny. Fig. 7. 

Recorded from northern South West Africa and Zulu- 
land. Widespread but usually rather rare in Europe, Africa 
and America. 

S.W.A. — 1918 (Grootfontein): Grootfontein, Volk 1977. 

Natal. — 2832 (Mtubatuba): Hluhluwe Game Reserve, 
Ward 2574; 2575. 

Wild, in F.Z. 1: 378 (1961), records a possible second 
species ( Elatine ? ambigua Wight) from the Caprivi Strip, 
viz. Stephens 56 (BM), from pools close to the Linyanti 
River, Kazungula and Kasane. It differs from E. triandra in 
that it develops longer petioles (up to 2 mm long) and longer 
pedicels (up to 5 mm long), otherwise the two taxa are very 


by A. A. Obermeyer 

Perennial woody shrubs or shrublets, often prostrate, rarely annuals, richly endowed with salt 
secreting glands. Leaves decussate or in verticils of 4, often in axillary fascicles through 
suppression of the branchlets. Flowers persistent, solitary or in some species aggregated apically in 
terminal dichasia with the subtending leaves sometimes reduced to bracts, sessile, regular, bisexual 
(polygamous and sometimes dioecious in Niederleinia). Calyx tubular, induplicate, 4— 6-lobed. 
Petals 4-6, free, imbricate, with a ligule attached to the claw on the inside (rarely absent), pink, 
mauve, red, white or yellow. Stamens 3-24 (mostly 6), free or filaments connivent and expanded 
below, filiform above, anthers bilocular, versatile, extrorse; staminodes present in Niederleinia. 
Ovary superior, 1-chambered, with 1-4 (usually 3) parietal or basal placentas bearing 1-numerous 
ovules attached to long funicles; style filiform, with as many branches as placentas, rarely capitate. 
Capsule enclosed in the calyx, dehiscing longitudinally; seeds small, ovoid-fusiform, smooth or 
rough, albuminous. 

A cosmopolitan family consisting of 5 genera (Hutchinson, Gen. FI. PI., 1967); Frankenia, the only genus represented 
in Southern Africa, being the largest and most widespread; the genera Hypericopsis, Beatsonia and Niederleinia 
monotypic, and Anthobryum with 5 species, all recorded from restricted, widely separated regions; all 4 genera closely 
related to Frankenia. Halophytes, usually gregarious, found along the coast and further inland, rarely in non-saline 

Literature of general interest: 

Marloth, R. Zur Bedeutung der salzabscheidenden Driisen der Tamariscinen. Ber. Dt. Bot. Ges. 5: 319-324 (1887). (Cf. 
p. 323 for reference to Frankenia ). 

Bray, W. L. The geographical distribution of Frankeniaceae in connection with their systematic relationships. Bot. Jahrb. 
24: 395-417 (1898). 

Gundersen. The Frankeniaceae as a link in the classification of Dicotyledons. Torreya 27: 65-71 (1927). 

Summerhayes, V. S. A revision of the Australian species of Frankenia. J. Linn. Soc. (Bot.) 48: 337-387 (1930). 

*The name was preserved for the family. 




Frankenia L., Sp. PL: 331 (1753); Gen. PI. ed. 5: 154(1754); DC., Prodr. 1: 349 ( 1824); Harv. in 
F.C. 1: 1 14 (1860); Oliv. in F.T.A. 1: 136 (1868); Niedenzu in Pflanzenfam. 3, 6: 286 (1895); ed. 
2, 21: 279 (1925); Adamson in FI. Cape Penins. 588 (1950); Phill., Gen. ed. 2: 508 ( 1951); Hutch., 
Gen. Flow. PI. 2: 321 (1967); Roessler & Schreiberin F.S.W.A. 91 (1968). Type species: F. laevis 

Nothria Berg., Descr. PI. Cap. 171 (1767). 

Low spreading woody shrublets, rarely annuals, often hairy and salt encrusted. Leaves 
decussate below, in verticils of 4 above, with leaf clusters (from suppressed branchlets) and flowers 
in their axils, sessile, subsessile or shortly petiolate, connected by an amplexicaul, usually ciliate, 
stipular ridge or sheath; lamina small, linear to ovate, flat or with the margins rolled back, often 
tightly so and thus terete or allantoid in shape, glabrous or pubescent, punctate. Flowers in forks of 
branches (determinate) sessile, small, persistent, scattered all over or aggregated in terminal 
dichasia (leaves subtending flowers in dichasia not differing from ordinary leaves in South African 
species). Calyx tubular, induplicate, 4— 5-fid with prominent ribs, glabrous, puberulous and 
occasionally with long scattered setae. Petals 4—5 (5 in South African species), pink, mauvy pink 
or white, with an adnate scale or ligule attached to the inside of the claw. Stamens 6 (rarely 4-5), 
filaments connivent and expanded below, filiform above, anthers bilocular, bipartite or nearly so, 
versatile. Ovary with 2-4 (3 in South African species) parietal placentas bearing one to many 
ovules; style terete, with 3 stigmatic lobes. Capsule ovoid to narrowly ovoid, seeds small, 
ellipsoid-fusiform, papillate or smooth, maturing very early. 

About 75 species (Hutchinson, 1967) recorded from the Mediterranean, along the Atlantic seaboard, western and 
southern Africa to western and southern Australia (the latter with about 45 species) to Asia and the Americas. Three species 
are found in Southern Africa, two endemic, the third almost cosmopolitan and weed-like in behaviour. Halophytes, found 
along the coast and further inland. 

The genus was named in honour of Prof. Franken or Frankenius, a professor at Uppsala, who died in 1661. 

Leaves ovate to lanceolate, flat or revolute, petiolate with the petiole 1 mm long; intemodes usually long; flowers very 
numerous, solitary or bunched at the nodes, about 5 mm long; seeds small, 0,5 mm long, smooth, yellow; annual 

or perennial; widespread 1 . F. pulverulenta 

Leaves allantoid or flat and linear, subsessile; intemodes usually short; flowers in apical dichasia or few, solitary and 
scattered, 9-11 mm long; seeds 1 mm long, rough, brown; woody perennials: 

Plants prostrate with short erect branchlets ending in many-flowered dichasia; stamens as long as petals, style longer, 

exserted; south-western Cape to eastern Cape (as far as East London) 2. F. repens 

Plants forming bushes up to 1 m tall; flowers few, scattered towards the tips of the branches; stamens shorter 

than petals; style just overtopping stamens; Namib 3. F. pomonensis 

1. Frankenia pulverulenta L., Sp. PI. 1: 

332 (1753); Harv. in F.C. 1: 115 (1860); Oliv. 
in F.T.A. 1: 136 (1868); Niedenzu in Pflanzen- 
fam. 3, 6: 287 (1895); ed. 2, 21: 279 (1925); 
Summerhayes in J. Linn. Soc. (Bot.) 48: 386 
(1930); Adamson in FI. Cape Penins. 589 
(1950). Type from Crete (LINN 457.6, holo.). 

F. nodiflora Lam., Encycl. 2: 396 (1786); 111. t.262, f.4 
(1792); DC., Prodr. 1: 349 (1824); Harv. in F.C. 1: 115 
(1860). Type; Cape of Good Hope (P, holo.). F. densa 
Pohnert in Mitt. Bot. Staatssamml. Miinchen 1: 447 (1954). 
Type: S.W. Africa, Grootfontein Slid, Dinter 8059 (M, 
holo.; PRE). 

Copiously flowering annuals or rarely pe- 
rennials with spreading, often reddish branches; 
intemodes usually long and bare, glabrous or 
puberulous with short curly hairs, branchlets 
often contracted, forming leafy fascicles at the 
nodes. Leaves with a short petiole (up to 1,5 
mm in the lower leaves), which is fused below 
to a stipular amplexicaul, ciliate (occasionally 
entire) sheath; lamina ovate, obovate, lanceo- 
late or oblanceolate, 3-5 (-7) mm long, flat or 
with the margin recurved to tightly revolute and 
then compressed-terete; upper surface glabrous, 
punctate, lower glabrous or puberulous. Flow- 



ers small, c. 5 mm long, very numerous, scat- 
tered and solitary in the forks of the branches 
(probably determinate) and flanked by side- 
branches, which are either well developed or 
repressed. Calyx tubular, 3,5-4, 5 mm long, ribs 
prominent, lobes very short and acute. Petals 
cuneate, 5 mm long, apex obtuse, erase, ligule 
longitudinally adnate to the claw, acute at the 
apex with free margins, pink, mauve or white. 
Stamens just exserted from the corolla-throat, 
didymous, the longest 2,5 mm long, filaments 
connivent, linear-lanceolate below, filiform in 
upper half; anthers small, somewhat bipartite, 
medifixed, semicircular. Ovary with many 
ovules (c. 45 ovules in South African plants), 
style with 3 terete, short stigmatic branches, just 
exserted above the stamens. Capsule with 
ovoid-fusiform, papillate, reticulate seeds about 
0,5 mm long, maturing early (possibly in a 
matter of days). 

A widespread species, probably dispersed by man and 
beast, occurring in Europe, Africa, Asia, Australia. Re- 
corded from S.W. Africa and along the Cape coast as far 
east as Port Elizabeth; further inland in the drier areas, in 
saline surroundings such as salt pans and banks of brackish 
streams. It flowers all the year round. Collections indicate 
that the plants may behave as annuals or occasionally persist 
as perennials when they become more woody and larger. 

S.W. A.— 2416 (Maltahohe): Grootfontein-Siid, possibly 
farm Lisbon, Dinter 8059. 2615 (Liideritz): Farm Sinclair 
Mine, Giess 2298; 2303. 2617 (Bethanie): Bethanien, 
Dinter 8270. 2618 (Keetmanshoop): River N. of Station 
Water, Merxmiiller & Giess 3606. 2619 (Aroab): 38 km 
W.N.W. of Aroab, Acocks 18072. 2717 (Chamaites): Great 
Fish River, Aiais, Oertendahl 635; Thorne sub SAM 52643. 
2818 (Warmbad): Bank of Ham River, Warmbad, Galpin 
s.n. 2819 (Ariamsvlei): Ariamsvlei, Walsersbrunn, Oerten- 
dahl 652. 

O.F.S. — 2726 (Odendaalsrus): 30 km W. of Odendaals- 
rus, Acocks 14010. 2926 (Bloemfontein): Haagenstad, 
Potts sub BLFU 6615. 

Cape. — 2818 (Warmbad): Henkries, Orange River, M. 
Schlechter 22. 2824 (Kimberley): Rooipan, c. 48 km 
W.S.W. of Kimberley, Leistner 1974. 2917 (Springbok): 
Nababeep, Hardy 1682. 2918 (Gamoep): Brakrivier, Pear- 
son 4868. 3118 (Vanrhynsdorp): 19 km S. of Bitterfontein, 
Taylor 5505. 3119 (Calvinia): Zwart Doom River W. of 
Brandkop, Compton 18894; 33 km E.N.E. of Loeriesfon- 
tein, Acocks 13202. 3124 (Victoria West): Hutchinson, 
Acocks 9589. 3218 (Clanwilliam): Piketberg, Compton 
15042. 3219 (Wuppertal): Beukesfontein, Pearson 5005. 
3224 (Graaff Reinet): near Kendrew P.O., Acocks 20616. 
3318 (Cape Town): About Salt River, Burchell 513; Green 
Point, Ecklon & Zeyher (Enum. 238); Robben Island, 
Walgate 613. 3320 (Montagu): Witteberge near Matjiesfon- 
tein, Marloth 11442. 3322 (Oudtshoom): Oudtshoom, The- 
ron 1038. 3325 (Port Elizabeth): Swartkops, Theron 1141. 
3421 (Riversdale): The Fisheries, Acocks 21566. 

2. Frankenia repens (Berg.) Fourc. in 
Mem. Bot. Surv. S. Afr. 20: 43 (1941). Type: 
Cape, Auge (SBT, holo.; PRE, photo.). 

Nothria repens Berg., Descr. PI. Cap. 171 (1767). Type 
as above. 

F. nothria Thunb., Prodr. 58 (1794); FI. Cap. ed. SchulL 
295 (1823). Type as for F. repens. F. krebsii Cham. & 
Schlechtd. in Linnaea 1: 36 (1826). Type: Cape of Good 
Hope, Krebs (Bf). F. capitata and varieties, sensu Harv. in 
F.C. 1: 114 (1860). F. hirsuta sensu L., quoad coll. 
Tulbagh No. 195; cf. Jackson, Suppl. Proc. Linn. Soc. 
Session 130: 11 (1917-18); sensu Marloth, FI. S. Afr. 2: 
185, f. 124 (1925); sensu Adamson in FI. Cape Penins. 589 
(1950); non L. (1753). 

Prostrate woody shrublets, the spreading 
branches with numerous short upright branch- 
lets usually ending in flowering dichasia, 
glabrous or roughly curly puberulous. Leaves 
subsessile, allantoid, 4—7 mm long, rarely flat 
and linear, dorsally smooth, punctate, ventrally 
with white papillae or minutely puberulent and 
punctate. Flowers in dense to few-flowered 
dichasia, congested near the apices of the branch- 
lets. Calyx 7 mm long, often with long white 
coarse hairs on the ribs and minutely puberulent 
in the grooves. Petals obovate, c. 1 1 mm long, 
erase, pink to mauvy pink. Stamens with fila- 
ments 1 1 mm long, somewhat expanded in 
lower half, filiform above. Ovary cylindrical, 
with the style and 3 stigmatic branches exserted 
for 2-3 mm beyond stamens; ovules 21-35. 
Capsule with papillate, brown, broadly 
fusiform seeds c. 1,3 mm long. Fig. 8. 

Endemic and fairly frequent along the coasts of the 
Cape from Namaqualand to Port Elizabeth, on sea-facing 
rock ledges, salt pans, saline flats and lagoons. Further 
inland it is recorded from brackish streambanks, edges of 
saltpans and calcareous sandveld. “Lightly browsed by 
cattle and sheep” fide P.A.B. van Breda. 

A glabrous form is met with from Riversdale to Port 
Elizabeth, accompanied however by the more common 
pubescent form. It flowers during the summer months, from 
December onwards. 

Cape. — 2818 (Warmbad): near Henkries, Phillips 1636. 
2916 (Port Nolloth): between Port Nolloth and Holgat 
River, Pillans 5193. 2917 (Springbok): Steinkopf, M. 
Schlechter 40. 3017 (Hondeklip Bay): Hondeklip Bay, 
Spitfire Rock, Pillans 17941. 3118 (Vanrhynsdorp): Olif- 
ants River mouth, Acocks 24164. 3218 (Clanwilliam): 
Clanwilliam, Leipoldt 4403; Elands Bay, Leach & Carp 
11358. 3317 (Saldanha): Danger Bay, Tolken 426; 9 km W. 
of Moorreesburg, Acocks 20663. 3318 (Cape Town): 
Langebaan, Saldanha Bay, Taylor 3774; Van Putten’s vlei, 
Compton 21878: 3319 (Worcester): Goudini, Van Breda 
21. 3325 (Port Elizabeth): at the mouths of the Coega and 
Swartkops Rivers, Zeyher 1955; near the Swartkops River, 
Zeyher 1060. 3418 (Simonstown): Noordhoek, Wasserfall 
679; Cape of Good Hope Nature Reserve, Batsata Cave, 
Taylor 6727. 3419 (Caledon): Bot River, Marloth 2786: 



Fig. 8. — 1, Frankenia repens, x 2'A; a, leaf (dorsal view) x 7; b, calyx, x 7; c, seed, x 7; d, petal, x 3 Vi; e, claw of petal 
with ligule, X 7; f, stamen, X 7 (Taylor 6727). 



Danger Point, Bond 768. 3421 (Riversdale): Still Bay, Muir 
20. 3422 (Mosselbay): Mosselbay, The Point, Britten 645; 
Great Brak River, Schlechter 5753. 

Common names: Soutrankie; Seaheath. 

This Cape species is closely related to F. hirsuta L. 
and Linnaeus placed a Cape specimen (No. 195) from a 
collection forwarded by the Governor Rijk Tulbagh under 
this Mediterranean species and reduced Nothria repens 
Berg, to synonymy. The flowers of the Cape species are, 
however, bigger and the stamens and style, with its 3 
stigmatic lobes, well exserted. It is closely related to F. 
pomonensis, endemic in the Namib, but the latter forms 
small bushes and bears fewer and smaller flowers. 

In Flora Capensis the species is named F. capitata 
Webb & Berth., a species described from the Canary Islands 
in Phyt. Canar. 1: 131 (1837). No mention is made in the 
original description of a ligule on the petals (which is 
common to most species), neither is it shown on the 
drawing (t. 16). The style moreover is depicted as shorter 
than the stamens. 

3. Frankenia pomonensis Pohnert in 
Mitt. Bot. Staatssamml. Miinchen 1: 446 
(1954); Roessler& Schreiber in F.S.W.A. 91: 1 
(1968). Type: S.W. Africa, Namib, Pomona, 
Dinter 6424 (M, holo.; PRE). 

Small perennial, woody, greyish brown 
shrublets, much branched and bushy, up to 1 m 
high, glabrous, densely leafy. Leaves subses- 
sile, allantoid, c. 3(-6) mm long, upper surface 
glabrous, lower minutely and densely puberul- 
ous or papillate. Flowers sparse, towards the 
ends of the branches. Calyx tubular, 5-6 mm 
long, glabrous. Petals obovate, c. 9 mm long, 
crinkly, margin obtuse, erose, white to deep 
lilac or dark pink. Stamens just exserted from 
the corolla throat, anthers small, bipartite, 
semicircular. Ovary narrowly ovoid, with about 
39 ovules, style with 3 short fairly thick stigmas 
a little longer than the stamens. Capsule with 
ovoid-fusiform, greyish brown, rough seeds 1 
mm long. 

Recorded only from the Namib in S.W. Africa, on 
brackish flats, dolomite outcrops facing the sea and border- 
ing lagoons. Flowering during summer. 

S.W.A. — 2615 (Liideritz): Liideritzbay, Marloth 4810b; 
Giess & van Vuuren 676; Elizabeth Bay, Merxmiiller & 
Giess 2353. 2715 (Bogenfels): Pomona, Dinter 6424; 
Bogenfels, Merxmiiller & Giess 2351; 6219; Dinter 4037. 


by A. A. Obermeyer 

Shrubs, trees or subherbaceous, often halophytic. Leaves exstipulate, sessile, alternate, 
usually scale-like, punctate with deep-seated salt secreting glands. Flowers usually very small, 
terminal, solitary, spicate, racemose or panicled, regular, hypogynous, bisexual, rarely unisexual 
and dioecious. Sepals usually 5-4, free or rarely connate below, imbricate. Petals 5-4, free or 
connate below, imbricate. Stamens numerous or 5, free or rarely connate, inserted on a glandular 
disk; anthers versatile, bilocular with longitudinal slits. Gynoecium one-chambered or imperfectly 
septate with 3-5 basal placentas; styles 3-5, free, or stigmas 3, sessile; ovules numerous — 2, erect, 
anatropous. Capsule splitting into 3-5 valves, coriaceous; seeds erect with an apical hairy tuft 
and/or central plumose awn; embryo straight, elongate, endosperm present or 0. 

A small family consisting of 4 genera and about 75 species, the genus Tamarix alone accounting for about 50 of these. 
Distributed mostly in the northern hemisphere and usually halophytic. 


Tamarix L., Sp. PI. 1: 270 (1753); Gen. PI. ed. 5: 131, No. 337 (1754); DC., Prodr. 3: 95 (1828); 
Harv. in F.C. 1: 119 (1860); Oliv. in F.T.A. 1: 151 (1868); Niedenzu in Pflanzenfam. 3,6: 293 
(1895), ed. 2, 21: 285 (1925); Phill., Gen. ed. 2: 508 (1951); Hutch., Gen. Flow. PI. 2: 323 (1967); 
Baum, Monogr. Tamarix, thesis, Hebrew University (1967). Podlech in F.S.W.A. 90: 1 (1968). 
Type species: lectotype, T. gallica L. Chromosome number: 2n = 24. 



Shrubs or trees with the leaves reduced to scales, sessile, alternate, occasionally decurrent 
and/or auriculate, amplexicaul or vaginate, glabrous, papillose or hairy, punctate with sunken, salt 
secreting glands. Inflorescence a simple or compound raceme on old and/or on young wood; bracts 
more or less resembling the leaf, 1 per flower. Flowers in soft, erect or drooping catkin-like 
racemes, shortly pedicelled, very small and very numerous, bisexual or rarely unisexual and 
dioecious. Sepals 4—5, free or connate at the base, glabrous or hairy. Petals 4-5, free, persistent or 
deciduous, longer than the sepals, white, pink or red. Disk saucer- to cup-shaped, often fleshy, dark 
coloured with the margin variously lobed. Stamens 4—10, fused basally with the disk; filaments 
free, thin; anthers apiculate or obtuse; in female flowers changed to staminodes. Ovary 
flask-shaped, one-chambered with 3(4-5) basal placentas bearing several to many ovules, styles 3, 
short, incurved. Capsule usually 3-valved, pyramidal, narrowed to the apex; seeds several to many, 
narrowly ovoid with an apical tuft of hairs and a plumose awn, endosperm 0. 

Species about 50, mainly distributed around the Mediterranean basin, westwards to England and the Canary Isles, 
eastwards as far as China; with one isolated species in south-western and southern Africa. Two species commonly cultivated 

in Southern Africa, T. gallica L., the French Tamarisk, and 

Tamarix usneoides E. Mey. ex Bunge, 
Tent. Gen. Tamaric. 74 (1852); Baum, Monogr. 
Tamarix, (thesis, Hebrew University, 
Jerusalem) 82, fig. 32 (1967), ined.; Podlech in 
F.S.W.A. 90: 1 (1968). Syntypes: Cape, Koup 
area between Blauwekrans and Bitterwater on 
the Gamka River, Dr'ege; Zwartruggens area, 
Zondags River, on the bank and in the stream, 
Dr'ege b (PRE!; K); Little Namaqualand, in the 
river near Natvoet and at the mouth of the 
Orange River, Dr 'ege; T’Kousie River and near 
Rietfontein in Namaqualand, Ecklon & Zeyher. 
Syntypes at LE; iso-syntypes in many herbaria. 

T. usneoides E. Mey. ex Drege, Zwei Pfl. Doc. 61, 63, 
92, 94, 225 (1838), nom. nud.; E. Mey. ex Niedenzu in 
Pflanzenfam. ed. 2, 21: 288 (1925), in clavis. T. articulata 
sensu Harv., F.C. 1: 119 (1860); sensu Oliv. in F T. A. 1: 
151 (1868), pro parte quoad specim. Welwitsch s.n. from 
Angola, non Vahl. T. austro-africana Schinz in Bull. Herb. 
Boiss. 2: 183 (1894). Syntypes: Cape, Drege, Ecklon 2130, 
Zeyher 725, Rehmann ; S.W. Africa, Wandres 19, Schenk 
253, 383, Schinz 406, Pohle, Luderitz 150, 32, Stapff. 
Hopfner 15, Fleck 743 (all at Z, presumably). T. angolensis 
Niedenzu in Engl., Pflanzenw. Afr. 3, 2: 531 (1921), in 
obs.; Pflanzenfam. ed. 2, 21: 288 (1925), in clavis.; Baum, 
l.c. 75, fig. 28 (1967). Type: Angola, Welwitsch 1086 
(sphalm. 86) (BM, lecto.; K). T. engleri Arendt, Beitr. 
Tamarix (thesis, Berlin University) 51 (1926), ined.; Baum, 
l.c. in synonymy. 

A gregarious, copiously branched shrub or 
tree up to 5 m high, with a deep taproot and 
spreading superficial roots from which adven- 
tive buds may give rise to new plants; bark 
brownish grey with rough transverse scars; 
upper branchlets congested, soft, pendulous, 
giving the tree a plumose appearance. Leaves 
reduced to sharp pointed, amplexicaul, vaginate 
scales, completely covering the young stems, 

T. ramosissima Ledeb. 

pale glaucous-green. Inflorescences massed 
apically, the upper parts of the branchlets turn- 
ing into simple many-flowered racemes; leaves 
changing into semi-amplexicaul, auriculate 
bracts with long patent apices, the flowers 
solitary in their axils. Flowers very small, 
shortly pedicellate, greyish or creamy white, 
unisexual and dioecious or rarely bisexual. 
Sepals 5, free, ovate, 1 mm, spreading. Petals 
5, free, oblong, 2 mm, obtuse, erectopatent. 
Stamens in male or bisexual flowers on long, 
thin filaments exceeding the petals, basally 
attached to a 5-lobed dark red, fleshy disc; 
anthers obtuse, pink; in the female flowers the 
flattened staminodes vary in shape, from nearly 
as long as the ovary to minute, the apex 
spathulate (if vestigial anthers are present) or 
apiculate. Ovary flask-shaped, 1,5 mm long, 
with 3 thick, short incurved stigmas; ovules 
several; in the male flowers the ovary reduced 
in size. Capsule pyramidal-attenuate, about 6 
mm long; seeds c. 3 mm, grain-like with a 
plumose apical awn and a tuft of hairs as long as 
the seed. Fig. 9. 

Recorded from the S.E. and N.W. Cape, South West 
Africa and Angola, frequenting dry sandy areas with subter- 
ranean, often saline water, such as salt pans and riverbanks; 
often the dominant tree of the community. 

S.W.A.— 1712 (Posto Velho): Kunene River at Otjinun- 
gua, Giess 8908; Story 5817. 1713 (Swartbooisdrift): 
Ososouu River at Otjiwero, De Winter & Leistner 5358. 
1914 (Kamanjab): Huab River, farm Alettasrust about 40 
km S. of Kamanjab on road to Fransfontein, De Winter 
3114. 2016 (Otjiwarongo): farm Onguati, Schwerdtfeger 
1/168. 2114 (Uis): Brandberg, Liebenberg 5009; 

Hochbrandberg, Numas-Binsenwiesen, BOss 1459. 2115 
(Karibib): Erongo Mts. near Usakos, Schlieben 10321. 



Fig. 9.— 1, Tamarix usneoides, branches, x 2VS>; a, branchlet with leaves and flower (one sepal removed), x 8; b, sepal, x 
25; c, petal, x 12; d, stamen, x 24; e, staminode, x 36; f, disc with bases of filaments, x 12; g, ovary, x 12; h, seed, 
x 9 (Hardy & Ihlenfeldt 1429). 



2214 (Swakopmund): Kuiseb River at Gobabeb, Hardy & 
Ihlenfeldt 1429 ($). 2216 (Otjimbingwe): Okapuka camp on 
farm Otjisewa, Klein Windhoek River, Wiss & Kinges 
718. 2314 (Sandwich Harbour): 10 km S. of Walvis Bay in 
Kuiseb River, De Winter 3179. 2417 (Mariental): near 
Mariental, Basson 221. 2517 (Gibeon): 40 km S.S.E. of 
Stampriet, Leistner 1815 V. 2617 (Bethanie): Leeu River 
near Seeheim, D'Ewes subNBG 84305; along tributary of 
Fish River, Bethanie, Gerstner 6287. 2816 (Oranjemund): 
Oranjemund, Merxmuller & Giess 2286; Arrisdrift, Schenk 
253. 2818 (Warmbad): Ramansdrift, Kruger 6. 2819 
(Ariamsvlei): Klein Karas, Groendoom, Oertendahl 363; 
Ariamsvlei, Walsersbrunn, Oertendahl 323. 

Cape. — 2816 (Oranjemund): Richtersveld, 

Sendlingsdrift, Orange River banks, Werdermann & Ober- 
dieck 592. 2817 (Vioolsdrif): Kuboos Mission, Marloth 
12331. 2819 (Ariamsvlei): Velloorsdrift, Van Son sub TRV 
31820. 2820 (Kakamas): near Augrabies Falls, Van Zinde- 
ren Bakker 1368; 1,5 km N. of Kakamas near Orange River, 
Leistner & Joynt 2820; Botha 2979. 2821 (Upington): 
between Upington and Keimoes, Theron 828 (¥). 3018 
(Kamiesberg): near Garies, Rodin 1374. 3019 (Loeriesfon- 
tein): farm Brackfontein near Loeriesfontein, Schlieben & 
Van Breda 9851. 3119 (Calvinia): Zwartdoom River W. of 
Brandkop, Compton 18895. 3222 (Oudtshoorn): Dwyka 
River, 16 km S.W. of Prince Albert Road, Theron 1316; 
Beaufort West, Marloth 2149. 3324 (Steytlerville): Groot- 
rivierpoort, Scharf 1364; 1365; 1366; 1369. 

Common names: Tamarix; Abikwa tree. 

Tamarix usneoides is a gregarious, sand-loving 
halophytic tree or shrub dependent on an ample supply of 
ground-water. In South Africa it is common along the banks 
of the Orange River from near Upington to its mouth and 
also occurs along its tributaries and around saline depres- 

sions in Namaqualand and further south-east. When pushed 
over by floods, the prostrate trunks send up straight young 
stems, while broken off branches may also take root where 
they are deposited further down stream. Zeyher collected it 
in the south-eastern Cape in 1826 in the Grootrivierpoort 
(3324 Steytlerville). The tree from which Zeyher collected 
his material could very likely be the solitary tree next to the 
old road at the drift, now a farm road (c/. notes by Scharf at 
PRE). It is still common along the rivers of that region. 

In South West Africa the species is quite common in 
the low-lying western desert areas, bordering sandy river- 
beds and brackish shallow pans. It stretches further north to 
southern Angola. Although it is usually unisexual and 
dioecious, an occasional plant in a colony will revert to the 
(ancestral?) bisexual form, when fertile stamens are pro- 
duced. Niedenzu separated these as T. angolensis. 

In and around Swakopmund the species hybridises 
with the introduced garden species T. ramosissima Ledeb., 
which elsewhere has also been found growing wild. It can 
be distinguished from the endemic species by its green 
colour and pink flowers; the leaves moreover are not 

T. usneoides is closely related to T. aphylla (L.) Karst, 
from north Africa and Arabia, which is also occasionally 
cultivated in South West Africa, but this species has smaller 
flowers which are always bisexual, the anthers apiculate, 
and the branches appear to be thinner and longer, resembl- 
ing Casuarina. 

The Cape specimens are frequently festooned with red 
galls and the flowers are often attacked by insects which 
damage the ovaries causing them to become sterile and 
globose in shape. 


by L. E. CODD 

Glabrous, often aromatic trees. Leaves alternate, simple, entire, gland-dotted, exstipulate, 
penninerved. Inflorescence axillary or terminal, cymose or flowers solitary. Flowers hermaphro- 
dite, regular. Sepals 3, imbricate, persistent. Petals 5-12, free or united into a tube at the base, 
rather thick, in 1 or 2 whorls. Stamens 6-12, hypogynous; filaments united in a tube; anthers adnate 
to the outer surface of the tube with longitudinal extrorse dehiscence. Ovary superior, 1 -locular; 
ovules 2-many on 2-6 parietal placentas. Style short, thick; stigmas 2-5. Fruit a berry. Seeds 
2-many, shining; endosperm oily and fleshy. 

A family of 5 genera and about 1 1 species, natives of tropical north and south America, east tropical Africa and 
Madagascar. The African genus Warburgia extends to Southern Africa. Several species are known to have aromatic or 
pungent bark which is used medicinally. 

The relationships of- the Canellaceae are probably with the woody Ranales (Magnoliales of Hutchinson), near to the 
Myristicaceae or Winteraceae. The structures described as 3 sepals above are, by some authors (Verdcourt, Hutchinson), 
regarded as 3 bracts, and the 2 rows of petals as calyx and corolla, respectively. On the other hand Wilson supports the 
interpretation given here. 




Warburgia Engl., Pflanzenw. Ost-Afr. C: 276 (1895); Burtt Davy, FI. Transv. 1: 218 (1926); 
Phill., Gen. e<± 2: 509 (1951); Verdcourt in F.T.E.A. Canellaceae: 1 (1956); Melchior & 
Schultze-Motel in Pflanzenfam. ed. 2, 17a II: 223 (1959); Hutchinson, Gen. Flow. PI. 1: 64 (1964); 
Wilson in Am. J. Bot. 53: 336 (1966); nom. conserv. (Taxon 17: 328, 1968). Type species: W. 
stuhlmannii Engl. 

Chibaca Bertol.f. in Mem. Accad. Sci. Istit. Bologna 4: 545 (1853). Type species: C. salutaris Bertol.f. 

Evergreen trees. Leaves dark green, shiny above. Inflorescence axillary, flowers solitary or in 
reduced 3 or 4-flowered cymes. Sepals 3, suborbicular. Petals 10, in 2 rows, the outer larger and 
thicker than the inner, pellucid gland-dotted. Stamens 10; staminal tube equalling or subequalling 
the petals; anthers sessile on the upper part of the tube, about as long as the staminal tube, linear, 
with a fleshy apical appendage. Ovary ellipsoid; ovules 10-30 in 1 or 2 rows on 5 placentas. Style 
enclosed in the staminal tube or just protruding, truncate at the apex, bearing 5 oval stigmatic 
patches round the sides. Fruit a globose or ovoid berry; pericarp coriaceous. 

An African genus of 3 species, 1 of which extends into 

Warburgia salutaris {Bertol.f. J Chiov. in 
Nuov. Giom. Bot. Ital. n.ser. 44: 680 (1937); 
Mendes in Bol. Soc. Brot. Ser. 2, 43: 338 
(1969); Ross, FI. Natal 247 (1972); Palmer & 
Pitman, Trees S. Afr. 3: 1523 (1972). Type: 
Mozambique, Fornasini s.n. (BOLOf). 

Chibaca salutaris Bertol.f. in Mem. Accad. Sci. Istit. 
Bologna 4: 545, t.23 (1853). 

Warburgia breyeri Pott in Ann. Transv. Mus. 6: 60 
(1918); Gerstner in Bantu Studies 12: 220 (1938); Doke & 
Vilakazi, Zulu-English Dictionary 20 (1948). Type: Trans- 
vaal, Letaba District, Makhutswi (“Macoutsie”) River, 
Breyer sub TRV 17573 (PRE, holo.!). 

Tree up to 10 m tall; bark rough and 
mottled, red on inner side; branches terete, 
striate, lenticellate. Leaves coriaceous, shiny 
above, paler below, densely pellucid-dotted, 
shortly petiolate; petiole 2-5 mm long, channel- 
led above; blade oblong to oblong-lanceolate or 
elliptic, 4,5-11 cm long, 1,5-2, 5 cm broad, 
veins obscure above, inconspicuous below; 
apex acute; base cuneate; margin slightly invo- 
lute. Flowers axillary, solitary or in abbreviated 
3-flowered cymes; peduncle 2-3 mm long; 
bracts 0,5 mm long, deciduous, leaving con- 
spicuous scars; pedicel up to 1,5 mm long. 
Sepals 3, suborbicular, 2 mm long, 3 mm broad, 
minutely ciliate. Petals in 2 whorls; outer 5 
subcoriaceous, obovate, concave, 4-5 mm long, 

3 mm broad; inner petals 5, yellow, thinner- 
textured, spathulate, 4 mm long, 1,5 mm broad. 
Staminal-tube 3—4 mm long, apex protruding 
slightly beyond the stamens; stamens 10, 1,5 
mm long. Ovary oblong-ovoid, 3 mm long, 1 
mm diam.; stigma subsessile, truncate, 

Southern Africa. 

obscurely 5-lobed; ovules about 15-20. Fruits 
ovoid to subglobose, 2-3 cm long, 1, 5-2,5 cm 
in diam.; pericarp coriaceous, wrinkled; seeds 
several, flattened. Fig. 10. 

A rare constituent of coastal forest in northern Zulu- 
land and of submontane forest in eastern and northern 

Transvaal. — 2229 (Waterpoort): Soutpansberg, Van 
Warmelo s.n. 2328 (Baltimore): Blouberg, Codd 8733; 
Strey & Schlieben 8611. 2330 (Tzaneen): near Tzaneen, 
McCallum sub PRE 15483. 2430 (Pilgrims Rest): Macout- 
sie River, Breyer sub TRV 17573; Mariepskop, Van der 
Schijff 5196; 5503; 5950; Govt. Forester s.n.; Blyde River 
Poort, Van der Schijff 4307. 

Natal. — 2731 (Louwsburg): Magut, Gerstner 3416. 
2732 (Ubombo): Lake Sibayi area, Tinley 197; False Bay, 
Gerstner 6974; Ward 3004. 2832 (Mtubatuba): Corridor, 
Umfolozi Game Reserve, Ward 3067. 

Known as isiBaha (Zulu), Shibaha (Tsonga) and 
Molaka (Sotho), the bark, especially the root bark, is 
pungent and bitter and is widely used as a medicine against 
colds and fever. Gerstner reports that this has led to regular 
cutting down of trees which may have an adverse effect on 
their regeneration. The leaves are also bitter and this helps 
in identifying the species in the field. 

As yet no flowering or fruiting material has been seen 
from Zululand but the specimens from this area agree well 
in vegetative characters with those occurring in the Trans- 
vaal. However, fertile material is important for a correct 
assessment to be made. 

Verdcourt, l.c., united the Transvaal W. breyeri Pott 
with the tropical African species, W. ugandensis Sprague, 
but the slight differences in texture, size and shape of the 
leaves suggest that this view may not be correct. The fruits 
of Transvaal specimens (possibly immature) also appear to 
be smaller and more wrinkled than those from Tropical 

The type of W. salutaris is from Mozambique and, 
according to Verdcourt in Regnum Vegetabile 40: 27 
(1965), is no longer extant. Good material from Mozam- 
bique is thus also required in order to establish its affinity. 
At present it seems to be related rather to the southern than 
the tropical populations. 

Fig. 10. — 1, Warburgia salutaris, flowering twig, x 1 (Strey & Schlieben 8611); a, fruit, X 1 (Van Warmelo s.n.); b, 
flower, x 4; c, staminal column, x 7; d, ovary, x 7; e, sepal, X 4; f, outer petal, X 4; g, inner petal, X 4 (Strey & 
Schlieben 8611). 




by A. A. Obermeyer 

Herbaceous perennials, rarely annuals, shrubs or trees. Leaves alternate, rarely opposite, 
simple, margin entire, dentate or incised; stipules 2, foliaceous, small or minute, persistent or 
caducous. Flowers solitary, axillary, or in compound, often reduced, inflorescences, bisexual or 
rarely polygamous, occasionally also with cleistogamous flowers, actinomorphic or usually 
zygomorphic. Sepals 5, usually free, persistent, quincuncial, sometimes produced beyond point of 
insertion. Petals 5, free, deciduous or persistent, imbricate or contorted, hypogynous; in 
zygomorphic flowers the anterior petal sometimes spurred. Stamens 5 (-15), usually hypogynous; 
anthers connivent in a ring around the ovary, bilocular, introrse, opening by longitudinal slits, 
thecae and connective often produced into dorsal and apical appendages; filaments 0 or short, 
simple or variously appendaged; the 2 anterior often forming caudate glandular spurs. Ovary 
superior, sessile, 1 -locular with 3 (-5) parietal placentas bearing l-°° ovules; styles completely 
united, straight, oblique or sigmoid, often somewhat swollen in upper half, stigmas usually entire, 
apical. Capsule loculicidal often with contractile valves; rarely a berry or a nut. Seed with a short 
funicle, sometimes winged, tomentose or smooth, sometimes hygroscopic becoming glutinous 
when wetted, occasionally with a gibbous aril, with abundant endosperm. 

Genera 20; species about 800. Mostly pantropical with the large cosmopolitan genus Viola predominantly temperate. 

Flowers actinomorphic; trees or shrubs 1. Rinorea 

Flowers zygomorphic, the anterior petal longer or shorter, often spurred; herbs or under- 

Pedicels articulate; stipules entire, small, linear- acuminate; anterior petal much longer than 

the other 4; with or without a spur 2. Hybanthus 

Pedicels inarticulate; stipules small, with 2 basal teeth or foliaceous, laciniate; 

anterior petal shorter than the other 4, spurred 3. Viola 

5262 1. RINOREA 

Rinorea A ubl., Hist. PI. Guian. Franc. 1: 235, t. 93 (1775); Melchior in Planzenfam. ed. 2, 21: 349 
(1925); Phill., Gen. ed. 2: 510 (1951); Robson in F.Z. 1: 246 (1960); Hutch., Gen. Flow. PI. 2: 330 
(1967). Type species: R. guianensis Aubl. 

Alsodeia Thouars, Hist. Veg. Rec. 55 (1806); Oliv. in F.T.A. 1: 106 (1868). 

Trees or shrubs. Leaves alternate, rarely opposite or whorled, petiolate, simple, serrate, rarely 
entire, with reticulate, arched, closed veins; stipules small, covering very young, pointed leaf-buds, 
caducous. Inflorescence simple or compound, axillary and/or terminal; pedicels articulated; bracts 
small, caducous. Flowers actinomorphic, small. Sepals 5, quincuncial, persistent, short and hard. 
Petals equal or subequal, soft, erect or reflexed, white, greenish white, yellow or red. Stamens 5, 
the filaments free or connate below, with or without appendages; anthers with the connective 
variously appendaged and occasionally also bearing thecal appendages. Ovary with 3 parietal 
placentas, style straight, stigma terminal, small, ovules 1-several on each placenta. Capsule 
3-valved, the inner tissues woody, tearing apart from the outer skin, rarely semi-succulent or a 
berry. Seed 1-3 per valve, compressed, hard, shiny. 



Fig. 11. — 1, Rinorea angustifolia, flowering twig, X 1; a, longitudinal section of flower, x 4; b, stamen, x 7; c, capsule 
and seed, x 2; d, bud, X 4 (Guy 129). 



Species about 250, tropics and subtropics; 2 species in Southern Africa. 

Leaves 4-8 cm long, soft, dark green, closely dentate, on slender, shortly hairy dark branchlets; leaf scars not 

conspicuous; petals spreading to recurved 1. R. angustifolius 

Leaves 8-14 (-22) cm long, leathery, glaucous, coarsely spinoso-dentate, on stout pale green glabrous branches; 

transverse leaf scars prominent; petals erect to spreading 2. R. ilicifolius 

1. Rinorea angustifolia (Thouars) Baill. 
in Bull. Soc. Linn., Paris 1: 582(1886); Kuntze, 
Rev. Gen. PI. 1; 42 (1891); Melchior in Pflan- 
zenfam. 2, 21: 352 (1925); Perr. in Mem. Inst. 
Sci. Madag., ser B, 2, 1949: 328 (1950); in 
Humbert, FI. Madag. Fam. 139, Violacees: 36 
(1955); Tennant in Kew Bull. 16: 412 (1963). 
Type: Madagascar, without precise locality, Du 
Petit Thouars s.n. (P, holo.). 

Alsodeia angustifolia Thouars, Hist. Veg. Rec. ed. 2: 57, 
1. 18, fig. 1 (1806); DC., Prodr. 1: 313 (1824). Type as 
above. A. ardisiiflora Welw. ex Oliv. in F.T.A. 1: 108 
(1868). Type: Angola, Cuanza Norte, Pungo Andongo, 
Mata de Pungo, Welwitsch 885 (LISU, lecto; BM; K). A. 
natalensis (Engl.) Bak.f. in J. Linn. Soc. (Bot.) 40: 22 
(1911). Syntypes: Natal, Inanda, Wood 1001 (Bf; BM; 
GRA; K; NH; SAM); Pondoland, Intsubana, Bachmann 
1005 (Bf). 

Rinorea natalensis Engl, in Bot. Jahrb. 33: 135 (1902); 
Brandt in Bot. Jahrb. 50, Suppl.: 411 (1914); Bews, FI. 
Natal & Zululand 140 (1921); Melchior in Pflanzenfam. ed. 
2, 21: 350 (1925); Henkel, Woody PI. Natal 84 (1934). 
Type as above. R. ardisiiflora (Welw. ex Oliv.) Kuntze, 
Rev. Gen. PI. 1: 42 (1891); Sim, For. FI. Cape Col. 125, 
1. 15, 1 (1907); For. FI. P.E. Afr. 1 1 (1909); Bews, FI. Natal 
& Zululand 140 (1921). Type as above. 

Evergreen trees or shrubs up to 10 m high 
with a smooth, mottled grey bark, the juvenile 
parts and inflorescences puberulous. Branches 
slender, terete, brown. Leaves alternate, widely 
spaced, elliptic, 5-9 cm long, 2-4 cm broad, 
base cuneate, apex acuminate, margin serrate, 
the teeth gland-tipped, soft, dark glossy green, 
paler below; petiole short, stipules small, 
caducous. Inflorescences axillary, short, sub- 
corymbose, several- 1 -flowered; peduncle and 
pedicels slender, bracts minute. Flowers 
sweetly scented, small; buds conical, reddish. 
Sepals ovate, 2 mm, persistent. Petals oblong- 
elliptic, c. 7 mm long, upper half reflexed, 
glabrous or velvety towards the base, white. 
Stamens with the filaments fused below and 
with a dorsal appendage forming a short fringed 
or irregularly lobed cup, or appendage absent 
(in specimens seen from Zululand and Trans- 
vaal); anthers with an apical, erect, exserted, 
ovate-acuminate, petaloid, reddish-purple 
connective-appendage, thecae with an apical, 
small Ungulate appendage or this reduced to a 
mucro or 0. Ovary globose, glabrous (in S. Afr. 
specimens examined), style erect, terete, ex- 

serted, c. 4 mm long; stigma minute, apical. 
Capsule ellipsoid, c. 1,3 cm long, obtusely 
3-lobed, hard, muricate, glandular-punctate; 
seed 1 (-2) per valve, compressed-ovoid, 5 mm, 
smooth, pale yellow. Fig. 12. 

Common as an understorey tree (rarely up to 10 m tall) 
or shrub in evergreen coastal forests of Pondoland, Natal, 
Zululand and north eastern Transvaal; tropical Africa and 
Madagascar. Flowering August to October; occasionally 
earlier or later. 

Transvaal. — 2230 (Messina): Soutpansberg, Entabeni 
Forest Reserve, Obermeyer sub TRV 30275. 2330 (Tza- 
neen): Westfalia Estate, Scheepers 766; Woodbush Forest 
Reserve, De Hoek, Botha sub PRF 7411; Magoebaskloof, 
Gerstner 5862. 2531 (Komatipoort): road from Barberton to 
Piggs Peak, West 3013. 

Natal. — 2731 (Louwsburg): Ngome Forest Reserve, 
Gerstner 5113; Sim 2918; Sokosoko Forest, Gerstner 4911. 
2732 (Ubombo): Gwalaweni Forest, Moll 4451. 2830 
(Dundee): Qudeni Forest Reserve, Strey 9317; Wood 7717. 
2831 (Nkandla): Nkandla Forest, Acocks 11799; Gerstner 
3583; Ngoye Forest, Nicholson 434. 2832 (Mtubatuba): 
Hluhluwe Game Reserve, Ward 1426. 2930 (Pieter- 
maritzburg): Karkloof, Wylie sub Wood 10046; Hilton 
Road, Wood 10081. 2931 (Stanger): Durban, Oliver 6. 
3030 (Port Shepstone): Umtamvuna River, Strey 5834; 
Paddock, Strey 5997. 

Cape. — 3129 (Port St Johns): Egossa, Sim 2419; 2439; 
Mkambati Leper Institute, Marais 1192; Port St Johns, 
Hafstrom & Acocks 1932. 3328 (Butterworth): The Haven, 
Gordon-Gray 1340. 

In collections from the Gwalaweni Forest and the 
Transvaal the stamens have short free filaments; the dorsal 
appendage which fuses the filaments into a cup with a 
free ciliate margin which is found in typical material, is 
apparently suppressed. 

2. Rinorea ilicifolia (Welw. ex Oliv.) 
Kuntze, Rev. Gen. PI. 1: 42 (1891); Brandt in 
Bot. Jahrb. 50, Suppl.: 412 (1914); Keay in 
F.W.T.A. ed 2, 1: 101 (1954); Robson in F.Z. 
1: 250 (1960); Tennant in Kew Bull. 16: 409 
(1963). Type: Angola, Cuanza Norte, Pungo 
Andongo, Barrancos de Catete. Welwitsch 889 
(LISU, lecto; BM; PRE). 

Alsodeia ilicifolia Welw. ex Oliv. in F.T.A. 1: 108 
(1868). Type as above. 

Small understorey evergreen tree or shrub 
up to 5 m high. Branches terete, stout, pale 
green with dark transverse scars of the caducous 
stipules. Leaves alternate, coriaceous, glauc- 
ous, paler below, elliptic, usually about 8-16 



Fig. 12. — 1, Rinorea ilicifolia, flowering twig, X 1; a, petal, X 7; b, stamen, x 7; c, sepal, x 9; d, capsule and seeds, x %; 
e, flower, x 5 (Tinley 474). 



cm long and 4—8 cm broad, margin serrate to 
spinoso-dentate; petiole 1-2 cm, stout. Flowers 
very small, arranged in compound, terminal or 
axillary, elongated panicles which are shorter 
than the leaves, bearing small clusters of brown 
velvet buds, bracts deciduous, small; pedicels 
short. Sepals broadly ovate, 2-3 mm long, 
obtuse, finely ribbed fan-wise with a membran- 
ous margin, persistent. Petals erect, narrowly 
elliptic, about 5 mm long, yellow, deciduous. 
Stamens on very short filaments, fused at the 
base; apical connective-appendage ovate- 

acuminate, petaloid; thecal appendage small to 
minute. Ovary globose, glabrous, style erect, 
terete, exserted, stigma apical, minute. Capsule 
globose, obtusely 3-lobed, about 1-2 cm in 
diam., hard, brown, verrucose; seeds angled, 
about 6 mm long, yellowish. Fig. 12. 

Recorded from tropical and subtropical Africa as far 
south as Zululand, in damp evergreen coastal forests, where 
it may be the dominant understorey shrub. Flowering 
August to October. 

Natal. — 2732 (Ubombo): Gwalaweni Forest, Moll & 
Nel 5536; Tinley 474; Dutton & Tinley 21. 2832 
(Mtubatuba): Hluhluwe Game Reserve, Ward 2313. 

5271 2. HYBANTHUS 

Hybanthus Jacq., Enum. Syst. PI. Ins. Carib. 2 (1760), nom. conserv.; Melchior in Pflanzenfam. 
ed. 2, 21: 357 (1925); Phill., Gen. ed. 2: 510 (1951); Robson in F.Z. 1: 254 (1960); Roessler in 
F.S.W.A. 87: 2 (1969). Type species: H. havanensis Jacq. 

Ionidium Vent., Jard. Malm. 1: sub t.27 (1803); Sond. in F.C. 1: 74 (1860). 

Annuals, perennials, shrublets or shrubs. Leaves alternate and/or opposite, elliptic, ovate or 
linear, sessile or petiolate; stipules small, subulate or ovate, usually persistent, margin serrate to 
subentire, the teeth gland-tipped. Flowers solitary, axillary, rarely massed in terminal inflores- 
cences, pendulous, persistent; pedicels articulated, the 2 bracteoles opposite or subopposite at or 
below articulation. Sepals free, more or less equal, not produced below. Petals unequal, the 
anterior (in S. Afr. species) much larger than other 4, with or without a basal pouch, clawed in the 
middle, expanded above into a spathulate or broad lamina; the other 4 somewhat longer than the 
sepals. Stamens 5, free or connate at the base, similar or the 2 (4) anterior each with a recurved 
caudate nectariferous spur projected into the pouch of the anterior petal; anthers connivent with the 
connective produced above, petaloid. Ovary with 3 parietal placentas, 3-many ovules; style terete, 
short or long, straight or curved. Capsule loculicidal with 3 elastic valves; seed broadly fusiform or 
ovoid-globose, apex discoid, raphe distinct or indistinct, aril usually prominent; testa striate, 
punctate or smooth, hygroscopic, swelling to form a glutinous covering, when wetted. 

Species about 100, distributed in tropical and subtropical regions, predominantly in South and Central America. Four 
species in southern Africa, 2 of these endemic. 

Common Names: Lady’s Slipper, Pink Lady’s Slipper. 

Anterior petal more than 3 mm long; flowers often showy, blue, pink or violet, rarely white; seed white or pale yellow, 
fusiform with a truncate, discoid apex, or globose: 

Anterior petal with a basal pouch protruding from between the anterior sepals; filaments of 2 anterior stamens 
villous, bearing a short recurved caudate appendage which is inserted in the pouch; capsule glabrous or 
scabridulous; seed fusiform: 

Perennial herbs or suffrutices (flowering in their first year), glabrous to pubescent, very variable in growth-form 

and leaf-shape; leaves and flowers not congested; capsule smooth, glabrous; widespread . .1. H. ermeaspermus 
Annual, small, erect, scabridulous herbs with long linear leaves and shortly pedicelled flowers 

congested on the short stems; capsule scabridulous; Kalahari region 2. H. densifolius 

Anterior petal without a basal pouch; filaments glabrous, entire, without an appendage; capsule glabrous or 

pubescent; seed globose, pitted 3. H. capensis 

Anterior petal 3 mm long; flowers very small, white; ripe seed black, smooth, shiny, globose to ovoid, 2 mm in diam., 

with a white protruding aril 4. H. parviflorus 



1. Hybanthus enneaspermus (L.) F. 
Muell., Fragm. 10: 81 (1876); Eyles in Trans. 
R. Soc. S. Afr. 5: 422 (1916), as H. ennea- 
spermum (Vent.) Torre & Harms; Engl, in Bot. 
Jahrb. 55: 398 (1919): Robson in Bol. Soc. 
Brot. Ser. 2, 32: 164 (1958); in F.Z. 1: 254 
(1960); Roessler in F.S.W.A. 87: 2 (1969). 
Tennant in Kew Bull. 16: 430 (1963). Type: 
Ceylon, Hermann 317 (BM). 

Viola enneasperrna L., Sp. PI. 2: 937 (1753). Type as 

Ionidium enneaspermum (L.) Vent., Jard. Malm. 1, sub 
t. 27 (1803); Oliv. in F.T.A. 1: 105 (1868). Type as above. /. 
thymifolium Presl, Bot. Bemerk. 11 (1844), nom. illegit., 
non Presl (1835); Sond. in F.C. 1: 74 (1860). Type: eastern 
Cape — Natal, “Omsamwubo, Omsamcaba, Omtendo, to- 
wards Port Natal, in grassy places”, Dr'ege (S; TCD). /. 
caffrum Sond. in Linnaea 23: 13 (1850); in F.C. 1: 74 
(1860). Type: Port Natal, Gueinzius 94 (S, holo; K; PRE, 
photo.). I. hirtum Klotzsch in Peters, Reise Mossamb. Bot. 
1: 148 (1861). Type: Mozambique, Rios de Sena, Peters 
(Bf). /. enneaspermum var. hirtum (Klotzsch) Oliv. in 
F.T.A. 1: 106 (1868), (sphalm. hirta). 

Hybanthus enneaspermus var. serratus Engl, in Bot. 
Jahrb. 55: 398 (1919); Burtt Davy, FI. Transv. 1: 130 
(1926). Syntypes: Transvaal, Pienaars River, Schlechter 
4218 (Bf; GRA; K; PRE); Mozambique, Ungulubi, 
Schlechter 12140 (Bt; BM; K; PRE). — var. caffer (Sond.) 
Robson in Bol. Soc. Brot. Ser. 2, 32: 169 (1958); in F.Z. 1: 
257 (1960). H. caffer (Sond.) Engl., l.c. 400 (1919). — var. 
angustifolius Engl., l.c. 400 (1919); Hulme, Wild Flow. 
Natal 1. 1, fig. 12 ( 1954). Type: No specimens cited. H. 
hirtus (Klotzsch) Engl., l.c. 399 (1919). — var. glabrescens 
Engl., l.c. 399 (1919); Burtt Davy, FI. Transv. 1: 131 
(1926). Syntypes: numerous collections from tropical Af- 
rica and inch Schlechter 11729 (Bf; GRA) from 
Komatipoort. H. thymifolius (Presl) Engl., l.c. 400 (1919). 

Perennial (flowering in first year) with the 
stems often deciduous, trailing or erect, clus- 
tered from a woody crown, 20-50 cm high, 
with few side branches. Stems terete or ridged, 
glabrous, scabrid or pubescent. Leaves alter- 
nate, very variable, linear to ovate, 1-9 cm 
long, 1-2,5 cm wide, apex acute to obtuse, base 
attenuate into a petiole or petiole 0, soft, glab- 
rous, puberulous to sericeous, margin serrate to 
more or less entire, the teeth gland-tipped (if 
teeth repressed, glands still persist); stipules 
small, subulate, 0,5-2 mm, gland-tipped. 
Flowers on long to short, thin pedicels; size of 
flower variable. Sepals free, subequal, linear- 
acuminate, to ovate-acuminate, 3-6 mm long, 
membranous, keeled, somewhat pubescent. 
Petals pink to bluish or violet; anterior much 
exceeding other 2 pairs in length, 5-20 mm 
long with a narrow claw about as long as the 
quadrate, emarginate lamina, at the base with a 
broad saccate bulge protruding from between 
the lower sepals; lateral petals asymmetrical. 

triangular, curved upwards, about half as long 
as anterior petal; posterior petals smaller and 
narrower. Stamens short, the filaments (and 
sometimes connective) of the 2 anterior tomen- 
tose, producing short caudate hairy appendages 
which descend into the pouch of the anterior 
petal; anthers connivent, connective produced 
into an ovate petaloid apical appendage. Ovary 
ovoid, style terete, stigma capitate. Capsule 
globose to ovoid, about 6-10 mm long, smooth 
with c. 3-4 seeds ripening on each placenta; 
seed fusiform to ovoid, apex and base discoid, 
c. 1,5 mm long, longitudinally ribbed, glabrous 
shiny, yellow to white, testa hygroscopic. Fig. 
13: 1. 

Widely distributed in Africa, Asia, Australia; wide- 
spread in South West Africa, Transvaal and Natal, often 
invading disturbed areas such as road verges, ploughed 
lands, riverbanks, forest margins, etc. 

S.W.A. — 1918 (Grootfontein): Aukas-Kreyfontein, 

Dinter 81 1 ; near Grootfontein, Schoenfelder S 385. 

Transvaal. — 2229 (Waterpoort): Waterpoort, Bre- 
mekamp & Schweickerdt 249. 2231 (Pafuri): Mazanji, 35 
km N.E. of Punda Milia, Codd & Dyer 4618. 2329 
(Pietersburg): near Mara, about 29 km from Louis Trichardt 
on Mara-Vivo road, Meeuse 10194. 2428 (Nylstroom): 
Mosdene, Galpin M576. 2430 (Pilgrims Rest): Shiluvane, 
Junod sub TRV 10205. 2431 (Acomhoek): Skukuza Camp, 
Acocks 16743. 2528 (Pretoria): Codrington, Thode A392. 
2529 (Witbank): 3 km N.E. of Middelburg, Meeuse 10263. 

Natal. — 2632 (Bela Vista): Ndumu, edge of Pongola 
flood plain, Pooley 111. 2832 (Mtubatuba): Hluhluwe 
Game Reserve, Ward 2403. 2930 (Pietermaritzburg): 
Pinetown district, Everton, Ward 5242; Isipingo, Wood 
12474. 2931 (Stanger): near Durban, Wood 1105. 3029 
(Kokstad): Port Shepstone, Umtamvuna Bridge, Strey 

Cape. — 3029 (Kokstad): 9 km from Umtamvuna Pont on 
Bizana Road, Acocks 10930. 3129 (Port St. Johns): Lom- 
bazi River Flats, Mogg 13563; Ntsubane, Strey 8562. 3130 
(Port Edward): 8 km inland from Umtamvuna River, Codd 

Common name: Pink Lady’s Slipper. 

Apparently the species is extremely variable and 
adaptable. The varieties caffer and serratus described for 
southern Africa and found elsewhere are here regarded as 
ecotypes. The flower, fruit and seed characters are stable 
throughout the range of the species but the leaf shows great 
variation, being narrow or broad, small or large, glabrous or 
pubescent, entire or serrate. 

2. Hybanthus densifolius Engl, in Bot. 
Jahrb. 55: 398 (1919); Melchior in Pflanzen- 
fam. ed. 2, 21: 359 (1925); Robson in F.Z. 1: 
258 (1960): Roessler in F.S.W.A. 87: 1 (1969). 
Type: S.W. Africa, Otjihua, Dinter 430 (Bf; K; 
SAM!; GRA!). 

Small, compact scabridulous annuals up to 
15 cm tall, the side branches mostly ab- 



Fig. 13. — 1, Hybanthus enneaspermus, x V 2 ; la, longitudinal section of flower, x 3 (Van der Schijff 1852). 2, H. 
capensis, leaf, x 1; 2a, flower, x 4; 2b, stamen, x 3 (Killick & Marais 2013). 2c, capsule, x 1; 2d, seed, x 2 
(Flanagan 2692). 3, H. densifolius, x 1; 3a, capsule, X 3 ( Dinter 6870). 4, H. parviflorus, X 1 (Hilliard 3147). 



breviated. Stems with short ridged intemodes. 
Leaves crowded, alternate, narrowly linear, 3-8 
cm long, 1-1,5 mm wide, apex acuminate, base 
attenuate into a thin petiole, margins inrolled 
with glands evenly spaced at intervals, forming 
shallow teeth. Flowers numerous, massed 
along the main stem in the leaf axils of the stem 
and abbreviated branches, on short pedicels, 
declinate, small. Sepals linear-lanceolate, 3 
mm, ciliate. Petals whitish, pink or purplish; 
anterior petal forming a basal pouch, clawed 
and with a spathulate lamina c. 7 mm long. 
Stamens with the 2 anterior tomentose, bearing 
a recurved spur. Ovary globose, scabridulous, 
style short, stigma obtriangular, pointing 
downwards. Capsule globose, c. 7 mm in 
diam., scabridulous; seeds stoutly fusiform, 
apex discoid, striate, yellowish. Fig. 13: 3. 

A small annual confined to the Kalahari region; South 
West Africa, Botswana, north-western Transvaal and north- 
ern Cape in sandy areas. A stable species, possibly self- 

S.W.A. — 1918 (Grootfontein): W. of Grootfontein, 
Schoenfelder S3 16. 2115 (Karibib): Karibib, Omaheke In- 
sel. Dimer 6973 . 2116 (Okahandja): Okahandja, Bradfield 
324. 2217 (Windhoek): Lichtenstein, Dimer 4354. 2316 
(Nauchas): Namib border towards Kuiseb, Strey 2598. 23 17 
(Rehoboth): Kalkrand on road to Rehoboth, De Winter 
3535. 2417 (Mariental): 10 km N. of Mariental, Acocks 

Transvaal. — 2229 (Waterpoort); Langjan Nature Re- 
serve, Zwanziger 742. 

Cape — 2624 (Vryburg): Armoedsvlakte, Henrici 36. 
2822 (Glen Lyon); 6 km W. of Padkloof Pass, Acocks 2062. 

3. Hybanthus capensis (Thunb.) Engl, in 
Bot. Jahrb. 55: 400 (1919); Robson in Bol. Soc. 
Brot. Ser. 2, 32: 165 (1958), in clavis; Batten & 
Bokelmann, Wild Flow. E. Cape Prov. 106, 
t. 86: 6 (1966), as H. capensis (Roem. & 
Schult.) Engl. Type: Cape, Uitenhage district, 
Galgebosch, Thunberg (LINN 1052.20). 

Viola capensis Thunb., Prodr. 40 (1794); FI. Cap. ed. 
Schult. 186 (1923). Type as above. 

Ionidium capense (Thunb.) Roem. & Schult., Syst. Veg. 
5; 393 (1819); Sond. in F.C. 1: 74 (1868). Type as above. I. 
natalense Harv. in F.C. 2: 585 (1862). Type: Port Natal, 
Sanderson 415 (K, holo.; PRE!, fragment of holotype). /. 
thorncroftii N.E. Br. in Kew Bull. 1921: 289 (1921). Type: 
Transvaal, Barberton, Reimers Creek, Thorncroft 1086 (K, 
holo.!; PRE, photo.). 

Hybanthus natalensis (Harv.) Burtt Davy, FI. Transv. 1: 
131' (1926); Robson in Bol. Soc. Brot. Ser. 2, 32: 165 
(1958), in clavis. H. thorncroftii (N.E. Br.) Burtt Davy in 
Kew Bull. 1924: 225 (1924); FI. Transv. 1: 131 (1926); 
Robson in Bol. Soc. Brot. Ser. 2, 32: 165 (1958), in clavis. 
H. sp. Hulme, Wild Flow. Natal t.41, f. 1 (1954). 

Small hispid suffrutices from a perennial 
rootstock. Stems terete, ribbed when young. 
Leaves alternate, ovate to obovate (1,5) 3(-7) 
cm long, (0,5) l(-3) cm wide, apex obtuse, base 
attenuate into a petiole, lower surface lighter 
coloured, margin shallowly crenate; stipules 
ovate-acuminate, membranous. Flowers decli- 
nate, on long thin pedicels, the sub-opposite 
bracts usually arising some distance below ar- 
ticulation. Sepals narrowly ovate-acuminate, 
fimbriate, the 2 lower slightly convex at the 
base. Petals white with lilac stripes, violet or 
mauve and a cream throat, the anterior 1,5 cm 
long, glabrous or tomentose near the base, 
without a pouch, claw short, lamina broadly 
spathulate; the other 4 somewhat longer than 
the sepals. Stamens short, with glabrous fila- 
ments, the connective-appendage oblong, ob- 
tuse to emarginate, orange. Ovary glabrous or 
pubescent, globose, style as long as ovary. 
Capsule pubescent or glabrous; seed obovoid 
with a small discoid apex and a distinct raphe 
and basal aril, smooth or faintly punctate. Fig. 
13: 2. 

Recorded from the Uitenhage district in the eastern 
Cape, northwards along the coast to Natal and the north- 
eastern Transvaal; in grassland (often ravaged by fires) or 
damp sheltered places, in valleys or margins of subtropical 
forests; rare. 

The collections from the Uitenhage district possess 
pubescent capsules. These plants appear to be depauperate, 
never exceeding 10 cm in height. The leaves are small and 
pubescent. It would seem to be an adaptation to more harsh 
surroundings. Several collections found in the more lush 
vegetation of the Transkei and further north are up to 40 cm 
tall and much bigger in all respects. 

Transvaal. — 2430 (Pilgrims Rest): The Downs, Rogers 
21878. 2531 (Komatipoort): Barberton, Williams sub TRV 

Natal. — 2731 (Louwsburg): Hlomohlomo Road near 
Ngotshe, Acocks 13051; Vryheid district, Ngwibi Mt., 
Hilliard & Burn 5907. 2831 (Nkandla): Eshowe, Gerstner 
2387. 2930 (Pietermaritzburg): Worlds View, Pieter- 
maritzburg, Fischer 909; Inanda, Wood 421. 2931 

(Stanger): Avoca, Schlechter 3006. 3030 (Port Shepstone): 
Campbellton, Dumisa, Rudatis 1744. 

Cape. — 3029 (Kokstad): Clydesdale, Tyson 1062. 3127 
(Lady Frere): Engcobo Mt., Flanagan 2692. 3128 (Umta- 
ta): Hill slopes at Baziya, Baur 290. 3228 (Butterworth): 
near Bashee River mouth, Acocks 12271 ; Kentani, Pegler 
157; The Haven, Gordon Gray 1135. 3325 (Port Elizabeth): 
Uitenhage, Prior s.n.; hill slopes at Algoa Bay, Zeyher 
1919. 3326 (Grahamstown): Howiesons Poort, Dyer 1725. 
3327 (Peddie): East London, Bokelmann 3; Cambridge, 
Wormald 41. 

4. Hybanthus parviflorus (L.f.) Baill., 
Bot. Med. 2: 481 (1884); Hilliard in Notes R. 
Bot. Gard. Edinb. 30: 128 (1970). Type: Cent- 
ral America, Mutis (LINN 1052.25). 

Viola parviflora L.f., Suppl. 396 (1782). Type as above. 



Perennial with prostrate woody branches. 
Stems with vertical lines of pubescence alternat- 
ing at the nodes. Leaves opposite or alternate, 
narrowly elliptic, 1-3 cm long, 5-9 mm broad, 
apex acute, base attenuate into a short to long 
petiole, margin serrate; stipules subulate. Flow- 
ers axillary, declinate, very small, c. 2-3 mm 
long, on thin pedicels up to 1 cm long in fruit. 
Sepals 2 mm long, curved upwards, pubescent. 
Petals white, short, the anterior 3 mm, with a 
broadly obovate, emarginate lamina. Stamens 
sessile, with an ovate, dark apical appendage. 

Ovary globose, with about 9 ovules; style short, 
oblique, stigma obtriangular. Capsule ovoid, 3 
mm in diam.; seed globose-ovoid, c. 2 mm in 
diam., black when mature, smooth, shiny, with 
a white spongy aril, hygroscopic. Fig. 13: 4. 

A native of South and Central America, apparently 
recently introduced. Recorded from several localities in 
Natal, usually in disturbed places. 

Natal. — 2829 (Harrismith): Oliviershoek Pass, High- 
land Sourveld, Acocks 23823. 2930 (Pietermaritzburg): 
Pietermaritzburg, Hilliard 3147; Barker 5128; Isipingo 
Flats, Ward 6634. 

5274 3. VIOLA 

Viola L., Sp. PI. 2: 933 (1753); Gen. PI. ed. 5: 402 (1754): Sond. in F.C. 1: 73 (1860); Becker in 
Pflanzenfam. ed. 2, 21: 363 (1925); Burtt Davy, FI. Transv. 1: 130 (1926); Phill. , Gen. ed. 2: 510 
(1951); Robson in F.Z. 1: 258 (1960); Hutch., Gen. FI. PI. 2: 334 (1967). Type species: Viola 
odorata L. 

Herbs, rarely shrublets, erect or trailing and rooting at the nodes. Leaves alternate, simple, 
petiolate, entire or usually serrate; stipules persistent, minute to foliaceous. Flowers usually single, 
axillary, declinate (cleistogamous flowers absent in African species); pedicels long, thin, 
inarticulate, bibracteate. Sepals 5, subequal, produced at the base below the point of insertion. 
Petals 5, unequal, obovate, the anterior spurred or saccate. Stamens with very short free filaments, 
anthers usually coherent in a ring around ovary, the 2 lower each bearing a recurved caudate 
nectariferous appendage protruded into the spur of the anterior petal; the anther connectives 
produced apically into a petaloid appendage. Ovary unilocular with 3 parietal placentas, 
multi-ovulate; style variously shaped, stigma terminal. Capsule 3-valved; seeds globose, pointed 
above and below, with a spongy aril. 

A very large cosmopolitan genus with probably about 300 species ( Hutchinson , 1967), mainly north temperate; V. 
odorata L. (Violet) and V. tricolor L. (Pansy or Heartsease) are popular garden plants and forms of the latter have been 
recorded as adventives in Southern Africa. Two closely related species occur in tropical Africa, one of these extending south 
to the Transvaal (Woodbush); one species with two varieties is endemic in the Cape Winter-rainfall Region. 

Herbaceous with creeping stems rooting at the nodes; leaves cordiform, soft, crenate; northern 

Transvaal 1. V. abyssinica 

Suffruticose with woody erect stems; leaves narrowly linear, somewhat succulent, hard; 

southern Cape 2. V. decumbens 

1. Viola abyssinica Steud. ex Oliv. in 
F.T.A. 1: 105 (1868); Becker in Pflanzenfam. 
ed. 2, 21: 364, 1. 159, fig. 34 (1925); Burtt Davy 
in FI. Transv. 1: 130 (1926); Keay in F.W.T.A. 
ed. 2, 1: 107, fig. 30 (1954); Robson in F.Z. 1: 
258, t.41 (1960). Type: Ethiopia, Schimper (K; 
PRE, photo.). 

Creeping or sometimes scrambling herbs 
from a perennial base, the long stems mostly 
unbranched, rooting at the nodes. Stems thin, 
narrowly winged. Leaves alternate, cordate, 
1-2 cm long and broad, crenate, hispid, lower 
side usually with scattered resinous bodies; 

petiole slender, up to 2 cm long, stipules 
foliaceous, laciniate, setulose. Flowers axillary, 
solitary, declinate, persistent; pedicels thin, up 
to 5 cm long, bibracteate, inarticulate. Sepals 
more or less equal, narrowly ovate-acuminate, 
c. 5 mm long, with rudimentary basal bulges 
produced below the point of insertion. Petals 
with the 2 upper pairs whitish, obovate, 5-10 
mm long; the anterior shorter, pale to dark blue 
or violet, veined, with a cylindrical large blunt 
spur adjoining the curved pedicel-apex. Sta- 
mens connivent, the apical connective appen- 
dage ovate, orange. Ovary globose, glabrous; 




Fig 14 —1 Viola abvssinica, x 1 (Bos 1 147); la, longitudinal section of flower, x 3; lb, capsule, x 1; lc, seed, x 1.2, 
l dei'umtn, .^decumtens, x 1 (Acock, 21M3 2a. MMM section of flower x 2, 2b capsule, x I 3. V. 
decumbens var. scrotiformis, x 1 (Thompson 192); 3a, longitudinal section of flower, x 2, 3b, capsule, 1. 



style narrowly funnel-shaped, oblique. Capsule 
ovoid, c. 5 mm long, glabrous, pale; seed ovoid, 
1-2 mm, several to many, with a distinct 
spongy aril. Fig. 14: 1. 

Widespread in tropical Africa. Recorded from Sout- 
pansberg and Woodbush forests, locally common in clear- 
ings and along paths. 

Transvaal. — 2230 (Messina): Soutpansberg, Entabeni 
Forest Reserve, Obermeyer sub TRV 30317. 2329 

(Pietersburg): Woodbush Forest Reserve, Hilliard 4753; 
Houtbosberg, Schlechler 4395. 2330 (Tzaneen): 

Duiwelskloof, Westfalia Estate, Bos 1147 ; Scheepers 843. 
2430 (Pilgrims Rest): The Downs, Junod 4260. 

2. Viola decumbens L.f., Suppl. 397 
(1782). Type: Locality and collector unknown 
(LINN 1052.19). 

Small perennial shrublet up to about 25 cm 
high with erect or spreading-erect, woody, mi- 
nutely papillate branched stems. Leaves alter- 
nate, linear, 15-25 mm long, 0,5-2 mm broad, 
apex acute, somewhat succulent, minutely 
papillate; stipules adnate to the sessile base of 
lamina, large and resembling the leaf or re- 
duced, with a small tooth on either side near the 
base. Flowers axillary, solitary, faintly scented, 
overtopping the leaves, on thin, usually long 
pedicels: bracts 2, subopposite in upper half or 
towards the apex, swollen below with 2 minute 
lateral teeth. Sepals narrowly ovate-acuminate, 
3-5 mm long, produced below point of inser- 
tion. Petals purple to violet, veined, the upper 2 
pairs subsimilar, narrowly oblong, 5—10 mm 
long; anterior shorter with a basal tubular blunt 
spur 2-5 mm long. Stamens with connivent 
sessile anthers, the apical connective appendage 
ovate, firm, orange; the 2 anterior stamens each 
with a long caudate nectariferous appendage 
projecting into the spur. Ovary globose with 
numerous ovules, style somewhat sigmoid. 
Capsule ovoid, 5-9 mm long; seed ovoid, 3 
mm, yellow, minutely punctate. 

Restricted to the southern Cape, from Worcester to the 
Cape Peninsula and eastwards as far as Riviersonderend 
Mountains; a montane species in fynbos, flowering July- 

The type of V. decumbens L.f. in the Linnaean 
Herbarium gives no information on locality or collector but 
it may be a duplicate of a Thunberg specimen which was 
collected in the Hottentots Holland Mountains. 

Two closely related varieties with a similar distribution 
are recognized. 

Leaves up to 50 mm long and 1 mm broad; flowers 
usually congested above; sepals 5-9 mm long; 
spur tubular, 3-5,5 mm long . . . .(a) var. decumbens 

Leaves about 20 mm long, 2-3 mm broad; flowers 
few, scattered along the branches; sepals 3,5-5 
mm long; spur saccate, 3 mm long and 
broad (b) var. scrotiformis 

(a) var. decumbens. 

V. decumbens L.f., Suppl. 397 (1782); Thunb., Prodr. 40 
(1794); DC., Prodr. 1: 299 (1824); Harv., Thes. Cap. 1: 30, 
t.46 (1859); Sond. in F.C. 1: 73 (1860); Marloth, FI. S. Afr. 
2: 189, t.64 d, fig. 126, B5— 6 (1925); Becker in Pflanzen- 
fam. ed. 2, 21: 373 (1925); Adamson in FI. Cape Penins. 
590 (1950); Jessop in Flow. PI. Afr. 38, 1. 1500 (1967). 
Type: Without locality or collector (LINN 1052. 19). — var. 
tenuis Bartl. in Linnaea 7: 540 (1832). Syntypes: Cape, 
Klein River Mountain, Ecklon & Zeyher (SAM); Zeyher 
1923 (PRE; S; TCD). 

Leaves linear, 3-5 cm long, 0,5-1 mm 
broad. Flowers aggregated at the top of the 
branchlets, exserted beyond the leaves. Sepals 
5-9 mm long. Spur tubular, blunt, up to 5,5 mm 
long. Capsule 9 mm long. Fig. 14: 2. 

Cape. — 3318 (Cape Town): Table Mountain, Schlechter 
12. 3418 ( Simonstown): Steenbras, Rogers 11025, Steen- 
bras Siding, Esterhuysen 2692; Hottentots Holland Moun- 
tains, Zeyher 1923; Betty’s Bay, White 5171 ; Pringle Bay, 
Compton 22721. 3419 (Caledon): Palmiet River Mouth, 
Acocks 21093; Hermanus, Fern Kloof, Anderson 8; Paar- 
deberg, Paviesvlei, Taylor 3603; Hangklip, Compton 
13502; Awila, E. of Witbaken, Thompson 1198. 

(b) var. scrotiformis (DC.) Jessop in 
Flow. PI. Afr. 38, 1. 1500 (1967). 

V. scrotiformis DC., Prodr. 1: 299 (1824); Sond. in F.C. 
1: 73 (1860); Becker in Pflanzenfam. ed. 2, 21: 373 (1925); 
Marloth, FI. S. Afr. 2: 189, t.64 c, fig. 126 A 1-4 (1925). 
Type: Cape Peninsula, Masson (BM, holo.; PRE, photo.). 
V. decumbens var. stipulacea Bartl. in Linnaea 7: 540 
(1832). Syntypes: Caledon, Swarteberg, kloof near the 
warm baths, Ecklon & Zeyher (SAM); Zeyher 1922 (PRE; 
S; TCD). 

Leaves shorter and broader than in the 
typical form, 2 cm long, 2 mm broad; flowers 
fewer, produced along the branches. Sepals 3-5 
mm long. Spur saccate, 3 mm long and broad. 
Capsule 5 mm long. Fig. 14: 3. 

Cape. — 3318 (Cape Town): Sneeukop near Wellington, 
Marloth 660; Stellenbosch, Dwarsberg, Jonkershoek, 
Thompson 191. 3319 (Worcester): Bainskloof, Schlechter 
1589; 9192; Compton 17492; Great Drakenstein Moun- 
tains, E. slopes of Bullers Kop, Esterhuysen 11873; 
Robertson district, Boesmans Kloof Pass at McGregor, 
Esterhuysen 5241. 3418 (Simonstown): Worcester district, 
Baviaans Kloof, Leighton 1355; Sir Lowry's Pass, Comp- 
ton 9004; Kogelberg Forest Reserve, near Rooiels River, 
Kruger 788. 3419 (Caledon): Elgin, Stokoe sub SAM 
63573; Zwarteberg among stones, Ecklon & Zeyher 1922; 
Galpin 3748; Schlechter 5537; Dasberg near Stormsvlei, 
Riviersonderend Mountains, Stokoe sub SAM 63577. 




(including Samydaceae) 

by D. J. B. Killick* 

Trees or shrubs, armed and unarmed. Leaves alternate, rarely opposite or whorled, simple, 
entire or toothed, teeth often glandular. Stipules present or 0. Inflorescence axillary or terminal, 
racemose, corymbose or paniculate, or flowers fasciculate or solitary. Flowers actinomorphic, 
bisexual or unisexual, monoecious or dioecious, occasionally polygamous. Sepals 3-6 or more, 
free or connate below into a calyx-tube, contorted or imbricate, rarely valvate, often persistent or 
accrescent. Petals as many as sepals or more or 0, usually small and inconspicuous, free, contorted 
or imbricate, with or without a scale on inner face. Disc extrastaminal or with free glands between 
stamens or of staminode-like scales inserted at base of petals, often adnate to the receptacle. 
Stamens 5-<», free or rarely connate, sometimes alternating with staminodes; anthers 2-thecous, 
dehiscing by longitudinal slits, rarely by apical pores. Ovary superior or more rarely semi-inferior, 
1-locular with 1 to many ovules on 2-8 parietal placentas, rarely ovules axile; styles 1-10, free or 
connate. Fruit a capsule or berry, 1 to many-seeded. Seeds usually small, sometimes arillate; testa 
woody, leathery or chartaceous; embryo straight; cotyledons usually broad; endosperm fleshy, 
copious, rarely thin or 0. 

Genera 89 with about 1250 species, mainly tropical and subtropical. 

Leaves opposite 6. Pseudoscolopia 

Leaves alternate: 

Petals with a fleshy gland on the inner face or staminodes petaloid with fleshy glands: 

Stamens numerous; petals spirally arranged 1 . Rawsonia 

Stamens 10-12; petals whorled 4. Kiggelaria 

Petals or petaloid staminodes without fleshy glands; petals sometimes absent: 

Petals present: 

Petals more than sepals: 

Branches spiny; fruit globose, indehiscent 2. Oncoba 

Branches without spines; fruit ovoid, beaked, dehiscing into 4—8 

valves 3. Xylotheca 

Petals as many as sepals: 

Stamens numerous, not collected in bundles opposite petals; fruit a 

berry 5. Scolopia 

Stamens equal in number to petals or in bundles opposite petals; fruit usually a 

Leaves with 5-9 nerves from base; flowers dioecious; seeds with a red 

aril 9. Trimeria 

Leaves penninerved; flowers bisexual; seeds without an aril: 

Style simple with a capitate or minutely bilobed stigma; flowers in pedunculate 

cymes; stipules deltoid 7. Gerrardina 

Styles 2-6 or 2-6-cleft; flowers in racemes or panicles; stipules absent or large 

and orbicular or reniform 8. Homalium 

*Except for Dovyalis by J. E. Langenegger. 



Petals absent: 

Leaves with linear or circular pellucid glands 13. Casearia 

Leaves without pellucid glands: 

Stamens in bundles of 3-10 alternating with disc glands 9. Trimeria 

Stamens not in bundles: 

Flowers bisexual; style very short; stigma peltate; leaves narrowly elliptic to 

oblanceolate 10. Aphloia 

Flowers unisexual or more rarely bisexual; styles 2-8; leaves elliptic to obovate 
or orbicular: 

Stamens surrounded by a ring of disc glands; ovary incompletely 4-8-locular 

with 2 ovules per loculus one above the other 11. Flacourtia 

Stamens intermingled with disc glands and alternating with them; ovules 1-6 

per placenta 12. Dovyalis 

5275 1 RAWSONIA 

Rawsonia Harv. & Sond. in F.C. 1: 67 (1860); Harv., Thes. Cap. 1: 20, t. 3 1 (1860); Sim, For. FI. 
P. E. Afr. 12 (1909); Engl., Pflanzenw. Afr. 3,2: 559 (1921); Gilg in Pflanzenfam. ed. 2, 21: 394 
(1925); Phill., Gen. ed. 2: 512 (1951); Wild in F.Z. 1: 262 (I960); Bamps in F.C.B. Flacourt. 4 
(1968). Type species: R. lucida Harv. & Sond. 

Evergreen shrubs or small trees. Leaves alternate, simple, shortly petioled, apparently 
stipulate (scar present); blade usually elliptic, coriaceous, margins spinulose-serrate. Flowers in 
axillary spike-like racemes, bisexual or male. Sepals 4—5, free, unequal, imbricate, concave. 
Petals similar to sepals but larger, with a petaloid scale opposite each petal with a gland on the 
inner face at the base. Stamens numerous, in several rows, the inner hypogynous, the outer 
attached to the base of the scales. Ovary superior, 1-locular, with a thick fleshy wall; ovules many 
on 4-5 parietal placentas; style almost none; stigmas 4—5. Fruit a globose berry. Seeds few, 
subglobose; testa leathery; embryo straight; cotyledons thin, elliptic; endosperm fleshy, copious. 

A genus of probably not more than 2 species occurring in southern and tropical Africa. The genus commemorates the 
Hon. Rawson W. Rawson, Secretary to the Cape Government and “ardent promoter and efficient patron of botany in South 

Rawsonia lucida Harv. & Sond. in F.C. 
1: 67 (1860); Harv., Thes. Cap. 1: 20, t.31 
(1860); Sim, For. FI. Cape Col. 128 (1907); 
Gilg in Bot. Jahrb. 40: 449 ( 1908); Sim, For. FI. 
P.E. Afr. 12 (1909); Bak.f. in J. Linn. Soc. 
(Bot.) 40: 23 (1911); Eyles in Trans. R. Soc. S. 
Afr. 5: 422 (1916); Engl., Pflanzenw. Afr. 3,2: 
559 (1921); Gilg in Pflanzenfam. ed. 2,21: 394 
(1925); Burtt Davy, FI. Transv. 1: 215 (1926); 
Henkel, Woody PL Natal 83 (1934); Wild in 
F.Z. 1: 262 (1960); Bamps in F.C.B. Flacourt. 4 
(1968). Type: Port Natal, Sanderson 118 (TCD, 
holo!; K!; PRE, photos). 

Tree up to 15 m high. Branches glabrous. 
Leaf-blade elliptic, occasionally obovate, 
10-15 cm long, 3-5,5 cm wide, apex acumi- 
nate, rarely acute, base narrowly cuneate to 

rounded, margins spinulose-serrate with in- 
curved teeth, midrib prominent on both sides, 
lateral veins 6-9, more prominent below, 
curved, looping near margin, rigidly coriace- 
ous, shiny, dark green above, paler below, 
glabrous; petiole 6-9 mm long, channelled, 
incurved. Flowers few; peduncles 1,2-2 cm 
long; pedicels 1-2 mm long. Sepals broadly 
ovate-triangular, 2-5 mm diam., concave, 
leathery, ciliolate along margin. Petals 3-5, 
unequal, broadly ovate-rotund, 3,5-8 mm long, 
4—7 mm wide, concave, ciliolate along margin, 
“deciduous”; petaloid scales elliptic- 
subrotund, 7 mm long, 4—5 mm wide, with 
somewhat fleshy tumid pubescent gland on 
inner face at base. Stamens: filaments 3-5 mm 
long; anthers narrowly ovate, slightly divaricate 
at base, 3,5 mm long. Ovary ellipsoid, 4-8 mm 



Fig. 15. — 1, Rawsonia lucida, twig with flower buds, x 1 (Kitlick & Strey 2466); a, bud, x 6 (PRF 3727); b, flower, x 6 
(PRF 3121); c, d, outer and inner sepals, x 6 (PRF 3727); e, petal, X 6 (PRF 3727); f, petaloid scale, X 6 (PRF 
3727); g, transverse section through ovary, X 6 (PRF 3727); h, fruit, X 1 (Story 5389). 



long, 4—5 mm wide, ridged; style almost none; 
stigmas 4-5, 1-1,5 mm long, spreading. Fruit 
yellow or green, 2,5-4 cm diam., with persis- 
tent style, “tardily dehiscent into 4-5 longitu- 
dinal sections when dry” (Wild, l.c.). Seeds 
few, subglobose, c. 1 cm diam. Fig. 15. 

A scrub-forest and forest tree occurring in the coastal 
areas of the eastern Cape and Natal, on the slopes of the 
Transvaal Drakensberg and in the northern Transvaal, 
Swaziland and tropical Africa. 

Transvaal. — 2230 (Messina): Makonde Mission, 9 km 
N.E. of Sibasa, Codd 6819. 2330 (Tzaneen): Woodbush, 
Hutchinson 2237. 2430 (Pilgrims Rest): Blyde River, 
Mariepskop Forest Station, Killick & Strey 2466 ; The 
Downs, Marais 131. 

Swaziland. — 2531 (Komatipoort): Kings Forest, Piggs 
Peak, Dlamini s.n. 2631 (Mbabane): Ubombo Mountains, 
2,4 km S. of Stegi, Keith s.n. 

Natal. — 2732 (Ubombo): Gwalaweni Forest, Moll 
4452. 2831 (Nkandla): Eshowe Forest, Codd 1857. 2832 
(Mtubatuba): Hluhluwe Game Reserve, Young 128. 2930 
(Pietermaritzburg): Krantzkloof Nature Reserve, Moll 
4033; Table Mountin, Killick 170; 687. 2931 (Stanger): 6 
km from Tongaat/Wartburg, Moll 919. 3030 (Port 

Shepstone): Elliott’s Farm, Paddock, Strey 7164. 

Cape. — 3129 (Port St. Johns): on way to Eagle’s Nest, 
Port St. Johns, Marais 1221 ; 1,8 km N of Umtata Mouth, 
Acocks 13573. 3228 (Butterworth): Kentani, Pegler 805 

With its rigidly coriaceous leaves with spinulose- 
serrate margins and incurved petioles, R. lucida is a 
clear-cut and easily recognizable species with several 
synonyms in tropical Africa. 

5284 2. ONCOBA 

Oncoba Forsk., FI. Aegypt.-Arab. CXIII, 103 (1775); Gilg in Bot. Jahrb. 40: 455 (1908); 
Pflanzenfam. ed. 2, 21: 402, 1. 172 (1925); Phill. , Gen. ed. 2: 512 (1951), pro parte; Wild in F.Z. 1, 
1: 275 (1960); Bamps in F.C.B. Flacourt. 16 (1968); Wild & Vidigal in FI. Mocamb. Flacourt. 15 
(1973). Type species: O. spinosa Forsk. 

Lundia Schumach. & Thonn., Beskr. Guin. PI. 231 (1827). 

Trees or shrubs, unarmed or with axillary spines; branches glabrous. Leaves alternate, simple, 
petiolate, stipulate; blade elliptic or ovate, margin serrate. Flowers bisexual, terminal or axillary, 
often on short sideshoots, usually large and showy. Sepals 3 or 4, free or united at base, imbricate. 
Petals 5-10 or more, exceeding the sepals, whorled. Stamens numerous, free; anthers linear. 
Ovary superior, 1-locular with 2-10 multiovulate parietal placentas; style simple. Fruit large, 
globose, with woody pericarp, smooth, indehiscent, many-seeded. Seeds with a leathery testa; 
embryo with leafy cotyledons. 

Forty species according to Flutchinson (1967), but possibly not more than 4 distributed in tropical Africa, southern 
Africa and Arabia. The generic name is derived from Onkob, the Arabic name for O. spinosa. 

Oncoba spinosa Forsk., FI. Aegypt.-Arab. 
CXIII, 103 (1775); Oliv. in F.T.A. 1: 115 
(1868); Gibbs in J. Linn. Soc. (Bot.) 37: 429 
(1906); Sim, For. FI. P.E. Afr. 12, t.2, fig. B 
(1909); Bak.f. in J. Linn. Soc. (Bot.) 40: 23 
(1911); R.E. Fr., Schwed. Rhod.-Kongo- 
Exped. 1; 155 (1914); Eyles in Trans. R. Soc. 
S. Afr. 5: 422 (1916); Gilg in Pflanzenfam. ed. 
2,21: 402, t. 172 (1925); Burtt Davy, FI. 
Transv. 1: 214, t.30 (1926); Hutch, in F.W.T.A. 
1: 161 (1927); Henkel, Woody PI. Natal 79, 91 
(1934); Exell & Mendonpa in C.F.A. 1, 1: 81 
(1937); Verdoom in Flow. PI. Afr. 28: 1. 1111 
(1950-51); Keay in F.W.T.A. ed. 2, 1: 188, t.71 
(1954); Wild in F.Z. 1, 1: 275 (1960); Bamps in 
F.C.B. Flacourt. 16 (1968); Wild & Vidigal in 
FI. Mocamb. Flacourt. 16 (1973). Type: 

Yemen, Hadie and Wadi Surdus, Forskal Herb. 
626 & 627 (C, syn.). 

Lundia monacantha Schumach. & Thonn., Beskr. Guin. 
PI. 231 (1827). Type: Ghana, Thonning 296 (missing). 

Oncoba monacantha (Schumach. & Thonn.) Steud., 
Nom. BoL ed. 2, part 2: 212 (1841). 

Shrub or small tree up to 5 m high. Bran- 
ches glabrous, conspicuously lenticillate with 
slender, sharp spines up to 5 cm long. Leaf- 
blade elliptic or ovate, 7-11 cm long, 3,5-5 cm 
wide, apex acuminate, base cuneate to rounded, 
margin serrulate or crenate-serrate, midrib and 
lateral veins sub-prominent above, prominent 
below, lateral veins 4—8 pairs, curved, glabrous, 
membranous to sub-coriaceous, dark green and 
shiny above, duller below; petiole 5-10 mm 



Pig. 16. — 1, Oncoba spinosa, flowering twig, x 1; a, fruit, x 1 (after Plate 1111 in Flowering Plants of Africa). 



long, channelled above. Flowers solitary and 
terminal or lateral on short axillary shoots, 
scented; pedicels 1,5-2, 7 cm long. Calyx 
globose in bud; sepals 4, joined at base, imbri- 
cate, elliptic, 1,5 — 1,8 mm long, c. 8 mm wide, 
strongly concave, green dorsally and white ven- 
trally. Petals white, 7-13, spreading, whorled, 
elliptic to obovate, the outer 3-4 mm long, 
1,3—1, 7 cm wide, slightly narrowed to a claw, 
the inner narrower. Stamens: filaments 4-8 mm 
long; anthers yellow, linear, c. 2 mm long with 
the connective produced into a triangular or 
minutely lobed tip or appendage. Ovary broadly 
obovoid to subglobose, 4,5 mm long, 3,5 mm 
wide, longitudinally sulcate; style c. 1 cm long; 
stigma patelliform, lobed at margin. Fruit yel- 
low, globose, c. 5 cm diam., smooth, marked 
with longitudinal lines, calyx and androecium 
persistent at base and gynoecium at apex, rind 
woody, hard, pulp yellow. Seeds numerous, 
ovoid-flattened, 6 mm long, 3 mm wide, testa 
shiny, brown. Fig. 16. 

A decorative shrub or small tree found on riverbanks, 
in woodland and scrub forest in South West Africa, the 
eastern Transvaal, Swaziland, Natal, tropical Africa and 

S.W.A. — 1725 (Livingstone): Mpilila Island, eastern Cap- 
rivi, Killick & Leistner 3397. 

Transvaal. — 2230 (Messina): Palmaryville, Codd & 
Dyer 4486; Sibasa, Van der Schijff 5255. 2330 (Tzaneen): 
Duiwelskloof, Scheepers 1113. 2430 (Pilgrims Rest): Calais, 
Renny 107. 2531 (Komatipoort): Barberton, Thorncroft 
HI 1255; Sabi River, E. of Skukuza, Wolhuter 4606. 

Swaziland. — 2631 (Mbabane): Lwandle, near Manzini, 
Miller S/176; Mbuluzi Poort, Stegi, Compton 31579. 

Natal. — 2632 (Bela Vista): Ndumu Game Reserve, Moll 
4235 A. 2831 (Nkandla): Eshowe, Lawn 2009 (NH). 2930 
(Pietermaritzburg): near Durban, Goldie sub Herb. Austro- 
Africanum 1702 (K, Z). 

Known as the Snuffbox Tree, Kafferklapper and um 
Thongwane (Z). The fruits are edible, but not palatable. The 
seeds yield a drying oil, but the mechanical separation of the 
seeds from the pulp makes this an uneconomical proposi- 
tion. The fruits are used to make snuff boxes, rattles for 
children and arm or ankle bands for dancers. The roots and 
leaves are said to be used medicinally by some tribes. The 
tree flowers between October and November. 

5284a 3. XYLOTHECA 

Xylotheca Hochst. in Flora 26: 69 (1843); Gilg in Bot. Jahrb. 40: 455 (1908); Pflanzenfam. ed. 2, 
21: 402 (1925); Wild in F.Z. 1,1: 272 (1960); Wild & Vidigal in FI. Mocamb. Flacourt. 1 1 (1973). 
Type species: X. kraussiana Hochst. 

Shrubs or trees, unarmed. Leaves alternate, simple, petiolate, stipulate; blade elliptic-obovate 
to broadly elliptic-obovate. Flowers solitary, male or bisexual, cymose or sub-umbellate in the 
upper leaf axils or terminal on the branchlets, large, sweet-scented. Calyx of usually 3 sepals, 
concave, free or almost free, imbricate, glabrous or pubescent. Petals 7-14, free or narrowed to 
base, imbricate. Stamens numerous; filaments free; anthers linear, dehiscing longitudinally from 
above. Ovary rudimentary in male flowers, 1-locular, multiovulate; ovules pendulous from c. 7 
parietal placentas; style terminal; stigmas equalling number of placentas, short, spreading. Fruit a 
woody capsule, splitting into c. 8 longitudinal valves; style persists as hard woody point. Seeds 
numerous, sometimes with a resinous aril. 

A tropical and southern African genus of about 10 species some of which are doubtfully distinct. The generic name 
refers to the woody pericarp of the fruit. 

Xylotheca kraussiana Hochst. in Flora 
26: 69 (1843); Gilg in Bot. Jahrb. 40: 455 
(1908); Pflanzenfam. ed. 2, 21: 402 (1925); 
Wild in F.Z. 1, 1: 274 (1960); Wild & Vidigal 
in FI. Mocamb. Flacourt. 12 (1973). Type: 
Natal, near Umlaas River, Krauss s.n. (352 in 

Herb. Kew, iso.!; PRE, photo.). 

Oncoba kraussiana (Hochst.) Planch, in Hook., Lond. J. 
BoL 6: 296 (1847); Harv. in F.C. 1: 66 (1860); Sim, For. FI. 
P.E. Afr. 12 (1909). O. tettensis sensu Hook.f. ex Harv. in 
F.C. 2: 584 (1862) quoad specim. Forbes (K!), excl. 
specim. Kirk (K!); Schinz in Mem. Herb. Boiss. 10:. 52 
(1900). O. macrofihylla sensu Schinz l.c. 



Fig. 17. — 1, Xylotheca kraussiana, flowering twig, X 1; a, pistil, X 3; b, transverse section of ovary, x 5; c, dehiscing 
fruit, x 1; d, arillate seed, x 1; e, fully dehisced fruit, x 'h (after Plate 1535 in Flowering Plants of Africa). 



Xylotheca lasiopetala Gilg in Bot. Jahrb. 40: 457 (1908); 
Pflanzenfam. ed. 2, 21: 402 (1925). Syntypes: Mozam- 
bique, Delagoa Bay, Monteiro 12 (K!); Junod (BR!; Z); 
Schlechter 11578 (COI; K!; L!); Quintas 68 (COI). X. 
kotzei Phill. in Kew Bull. 1922: 193 (1922). Type: Natal, 
Port Dumford Plantation, Kotze sub PRE 1479 (PRE, 
holo. !; K; sub PRF 3632). X. kraussiana Hochst. var. 
glabrifolia Wild in Bol. Soc. Brot. Ser. 2, 32: 53 (1958); in 
F.Z. 1, 1: 274 (1960); Killick in Flow. PI. Afr. 39, 1. 1535 
(1968); Wild & Vidigal in FI. Mocamb. Flacourt. 13 (1973). 
Type: Mozambique, Quissico, Exell, Mendonqa & Wild 
698 (BM, holo.!). 

Tree or shrub up to 10 m tall; branchlets 
glabrous or pubescent. Leaf-blade elliptic to 
broadly elliptic or obovate, 4—12 cm long, 
1,7 — 4 cm wide, apex acuminate, acute to 
rounded, base narrowly cuneate to rounded, 
margin entire, midrib prominent, veins 6-8 
pairs, dividing near margin to form conspicuous 
submarginal vein, membranous to chartaceous, 
glabrous or pubescent; petiole 0,5-1 cm long, 
glabrous or pubescent. Flowers in 1-3-flowered 
cymes in axils of leaves or on the branches; 
peduncles 2-3 cm long; pedicels up to 4,5 cm 
long, glabrous or pubescent. Sepals 3, ovate to 
obovate, 1,5- 1,8 cm long, 1 cm wide, deeply 
concave, apex rounded, green but margins 
white, pubescent outside. Petals white, 7-12, 
obovate, 2-3,5 cm long, 0,5-1, 5 cm wide, 
narrowed at base, glabrous or sparsely woolly. 
Stamens numerous; filaments 6 mm long, with 
scattered hairs; anthers linear, 5 mm long. 
Ovary ellipsoid, sulcate, densely hairy; style 

0,8- 1,0 cm long, columnar; stigmas 4, spread- 
ing, 1 mm long. Fruit woody, ovoid or ellipsoid 
with 8 longitudinal ridges, beaked, splitting into 
4—8 segments. Seeds numerous, reddish black, 
obovoid, 7 mm long, 5 mm wide, with fleshy 
red aril along one side. Fig. 17. 

Usually a forest margin shrub or tree found along the 
coast of Natal from the Port Shepstone area northwards into 
Mozambique. Also occurring in the north-eastern Trans- 

Transvaal. — 2231 (Pafuri): Nwambiya-sandveld, 
Brynard & Pienaar 4254; Van der Schijff & Marais 3684; 
Nyanda Bos, Van Wyk 4577. 

Natal. — 2632 (Bela Vista): Kwamuzimba, near Ndumu 
Game Reserve, Stephen 807. 2732 (Ubombo): 16 km 
Pongola Bridge/Maputa, Moll 4378. 2831 (Nkandla): 
Eshowe, Thode A 1223. 2832 (Mtubatuba): Enseleni River, 
Bayer 1456. 2930 (Pietermaritzburg): Shongweni Dam, 
Morris 805. 2931 (Stanger): Oqaqeni, Mapumulo, Edwards 
1791; 1708. 3030 (Port Shepstone): The Valleys, Mogg 
13841 ; Gibraltar, Strey 10565. 

A small tree or shrub with elliptic to broadly elliptic or 
obovate leaves, 4-12 cm long, with entire margins and a 
submarginal vein and large, showy flowers. Wild (l.c.) 
described var. glabrifolia and distinguished it from var. 
kraussiana by its glabrous vegetative parts. Examination of 
a wide range of material shows that this difference does not 
hold and the distribution is practically the same, therefore 
var. glabrifolia is not upheld in this treatment. It is 
interesting to note that A", kotzei, which Wild sinks under A. 
kraussiana var. kraussiana, has a holotype with sparsely 
pubescent vegetative parts, while the paratypes, apparently 
from the same tree, are completely glabrous. 


Kiggelaria L., Sp. PI. 2: 1037 (1753); Gen. PI. ed 5: 459 (1754); Harv. in F.C. 1: 7 (1860); Phill., 
Gen. ed. 2: 513 (1951); Wild in F.Z. 1; 265 (1960). Type species: K. africana L. 

Unarmed shrub or small tree. Leaves alternate, simple, petiolate, usually elliptic or ovoid, 
variously toothed, sometimes discolorous; stipules 0. Plants dioecious. Male flowers in axillary 
cymes, on longish pedicels. Female flowers solitary on long pedicels. Sepals 5-parted, segments 
narrowly elliptic, valvate, deciduous, puberulous. Petals 5, often very similar to sepals, elliptic, 
imbricate, each with a fleshy gland adnate at the base to the petals and free above, puberulous. 
Stamens 8-10, free; filaments short; anthers 2-thecous, opening by terminal pores. Ovary sessile, 
ellipsoid, 1-2-5 parietal placentas, densely pubescent; style short and thick, divided into 5 
branches. Fruit a globose, woody, several-seeded capsule; valves 2-5, pubescent, sometimes 
tuberculate. Seeds subglobose; testa somewhat woody; embryo straight; cotyledons flat, suborbicu- 
lar; endosperm fleshy. 

1 species (doubtfully more) occurring in Southern Africa and tropical Africa. The genus was named in honour of 
Francis Kiggelaer, a Dutch botanist. 



Fig. 18. — 1, Kiggelaria africana, twig with male flowers, X 1, (Thode A2603); a, male flower, X 3 (Thode A2603); b, 
female flower, X 3 (Fourcade 1591); c, fruit, X 1 (Mogg 2319); d, dehiscing fruit, x 1 (Mogg 2319). 



Kiggelaria africana L., Sp. PI. 2: 1037 
(1753); Pappe, Silv. Cap. 4 (1854); Harv. in 
F.C. 1: 71 (1860); Sim, For. FI. Cape Col. 128, 
t. iv (1907); Bak.f. in J. Linn. Soc. (Bot.) 40: 24 
(1911); Eyles in Trans. R. Soc. S. Afr. 5: 422 
(1916); Engl., Pflanzenw. Afr. 3,2: 571 (1921); 
Gilg in Pflanzenfam. ed. 2, 21: 413, 1. 179 (fig. 
f-h) (1925); Burtt Davy, FI. Transv. 1: 217 
(1926); Henkel, Woody PI. Natal 32, 73 (1934); 
Brenan, Checklist Tang. Terr. 233 (1949); 
Milne-Redh. in Mem. N.Y. Bot. Gdn. 8,3: 219 
(1953); Wild in F.Z. 1: 265 (1960). Type: 
“Aethiopia” (LINN 1191.3, holo. !) 

K. ferruginea Eckl. & Zeyh., Enum. 15 (1834—1835); 
Burtt Davy in Kew Bull. 1921: 337 (1921). Type: 
Namaqualand, Kamiesberg and near mouth of Orange 
River, Ecklon &Zeyher 118 (K, iso.!; PRE, photo.; B!; L!; 
P!; SAM!). K. integrifolia sensu Eckl. & Zeyh. l.c., non 
Jacq. (1788). K. dregeana Turcz., Animadv. 33 (1855). 
Type: Cape, “Zuurberg en by Bontjiesrivier”, Drege 
6722a (K, iso!; BM!; L!; P!; PRE!; S!; TCD!). — var. acuta 
Harv. in F.C. 1: 71 (1860). Type: without locality, Drege 
6722a (TCD, holo!; PRE, photo.). — var. obtusa Harv., l.c. 
Type: Cape, “sylvis in Sitzikamma (George), Olifantshoek 
(Uitenhage) et Kafferland”, Ecklon & Zeyher 117 (TCD, 
holo.!; PRE, photo.; K!). K. grandifolia Warb. in Engl., 
Pflanzenw. Ost-Afr. C: 278 (1895); Engl., Pflanzenw. Afr. 
3: 571 (1921). Gilg in Bot. Jahrb. 40: 468 (1902); Pflanzen- 
fam. ed. 2,4: 413 (1925); Type: Malawi, Buchanan 1469 
(B, holo.f; BM!; K!; PRE, photo.). K. africana L. var. 
obtusa (Harv.) Burtt Davy in Kew Bull. 1921: 336 (1921); 
FI. Transv. 1: 217 (1926). 

Shrub or tree up to 17 m high, semi- 
deciduous; bark pale grey, smooth; young 
branches stellately tomentulose, often rusty. 
Leaf -blade elliptic or ovoid, 4—11 cm long, 
1,5—4 cm wide, apex acuminate, acute or ob- 
tuse, apiculate, base cuneate to round, margin 
entire, serrulate, serrate or duplicate-serrate, 
midrib prominent below, lateral veins 5-9, 
prominent below, looping near margin, char- 
taceous to coriaceous, glabrous or white or 
rusty stellate, with tomentulose hairy pockets 
(acarodomatia) in axils of veins below, some- 
times discolorous; petiole 0,7-2, 5 cm long, 
channelled, glabrous or tomentulose. Male 
flowers yellowish-green, in 3-many-flowered 
axillary cymes; peduncle 0,3-2 cm long, tomen- 
tulose; pedicels slender, 0,5- 1,5 cm long, to- 
mentulose, minutely bracteate at base. Sepals 
narrowly elliptic, 3-4 mm long, 1,5 mm wide, 
apex acute, slightly keeled, stellate-pubescent. 
Petals elliptic, 4,25 mm long, 2,25 mm wide, 
apex obtuse to rounded, slightly keeled dorsal- 
ly, stellate, puberulous; basal scale oblong, 2 
mm long, 1,5 mm wide, toothed, fleshy. Sta- 
mens 10; filaments up to 1,7 mm long; anthers 

elliptic, 1,5 mm long, stellately puberulous. 
Female flowers yellowish-green, solitary in 
upper axils; pedicels slender, 1,7-2, 5 cm long, 
tomentulose. Sepals narrowly elliptic, 5,5 mm 
long, 1,5 mm wide, stellately puberulous. Pet- 
als elliptic 5,5-6 mm long, 2mm wide, stel- 
lately puberulous; scales ovoid 2-2,5 mm long, 
1-1,25 mm wide. Ovary ellipsoid, 2,5-3 mm 
long, 1,5-2 mm diam., densely pubescent; style 
short, divided into (4) 5 branches, c. 1,5 mm 
long, divergent, canaliculate, glabrous in upper 
part, minutely lobed. Fruit up to 1,5 cm diam., 
splitting from apex into 5 valves, yellowish- 
green, densely tomentulose, tuberculate. Seeds 
bright orange-red, 4—6 mm diam. Fig. 18. 

Occurring in all four provinces of the Republic in 
forest, usually fairly open forest, kloofs, along streams, in 
scrub and even in inhospitable places like the mountains of 
the Karoo. Also found in South West Africa, Rhodesia, 
Lesotho, Swaziland, Malawi, Mozambique and tropical east 

S.W.A. — 2317 (Rehoboth): Rehoboth, Strey 540. 

Transvaal. — 2328 (Baltimore): Blauwberg, Pole Evans 
893. 2329 (Petersburg): near Haenertsburg, Legat 133. 
2330 (Tzaneen): Modjadji’s Reserve, Krige 142. 2429 
(Zebediela): Schoonoord, Van Warmelo 25. 2430 (Pilgrims 
Rest): The Downs, Rogers 21936. 2527 (Rustenburg): 
Jacksonstuin, Van Vuuren 262. 2528 (Pretoria): 

Groenkloof, Mogg 14190. 2530 (Lydenburg): 6 km S.E. of 
Sewefontein, Codd 8101. 2531 (Komatipoort): Almeida, 
Nel 252. 2626 (Klerksdorp): Goedgedacht, Ventersdorp, 
Sutton 698. 2627 (Potchefstroom): Jack Scott Nature Re- 
serve, Wells 2444. 2628 (Johannesburg): Suikerbosrand, 
Prosser 1312. 2629 (Bethal): Ermelo, Leendertz sub TRV 
7784. 2630 (Carolina): 2 km E. of Forthill, Schlieben & De 
Winter 7980. 2730 (Vryheid): Oshoek, Devenish 176. 

O.F.S. — 2826 (Brandfort): Willem Pretorius Game Re- 
serve, Leistner 3017 . 2827 (Senekal): Doomkop, Goossens 
768; Braamhoek, Galpin 13938. 2828 (Bethlehem): 2,4 km 
S.E. of Bethlehem, Scheepers 1432. 2829 (Harrismith): 
Platberg, Puttrill s.n. 2926 (Bloemfontein): ThabaNchu, 
Roberts 1823A. 3027 (Lady Grey): Zastron, Heydorn 18. 

Swaziland. — 2631 (Mbabane): Hlatikulu Forest, Com- 
pton 28162; Forbes Reef Bush, Burtt Davy 2799 (BM). 

Natal. — 2730 (Vryheid): Tweekloof and Altemooi, 
Thode A180. 2731 (Louwsburg): Ngome Forest, Gerstner 
4491. 2829 (Harrismith): Cathedral Peak, Killick 986. 2830 
(Dundee): Mfongosi, Edwards 1290. 2831 (Nkandla): 
Nkandla Forest, Gerstner 3596 (NH). 2832 (Mtubatuba): 
Hluhluwe Game Reserve, Ward 2064. 2929 (Underberg): 
Bushmen’s Nek, Killick & Vahrmeijer 3977. 2930 (Pieter- 
maritzburg): Table Mountain, Killick 426. 3030 (Port 
Shepstone): Ifafa, Rudatis 390. 

Lesotho. — 2828 (Bethlehem): Leribe, Dieterlen 180. 
2927 (Maseru): near Cannibals Cave, Lekhatsi, Jacot Guil- 
larmod 843 ; Mamathes, Jacot Guillarmod 522. 3027 (Lady 
Grey): Thaba Tsueu, Page sub BOL 16790. 

Cape. — 3018 (Kamiesberg): Kamiesberg, Taylor 5538; 
“Kamiesberge and near mouth of Orange River”, Ecklon 
118 (B, K, L, P). 3026 (Aliwal North): near Burghersdorp, 



Flanagan 1550. 3125 (Steynsburg): Bo-Rietpoort, Van der 
Walt s.n. 3126 (Queenstown): mountain tops, Queenstown, 
Galpin 1564. 3127 (Lady Frere): Cala, Ferreira 2. 3128 
(Umtata): Pot River Berg, Maclear, Galpin 6577. 3129 
(Port St. Johns): Egossa Forest, Strey 8864; between 
Morley and Umtata River, Dr'ege 6722 b (K). 3218 (Clan- 
william): on banks of Olifants River, Galpin 11114. 3219 
(Wupperthal): Klein Koupoort Nek, Taylor 7481. 3222 
(Beaufort West): Nieuwveld Mountains, Marloth 8304. 
3225 (Somerset East): near Somerset East, Bolus 326 (K). 
3227 (Stutterheim): Komga, Flanagan 303; Happy Valley, 
Comins 1766. 3228 (Butterworth): Kentani, Pegler 738. 
3318 (Cape Town): Table Mountain, Flanagan 2464. 3319 
(Worcester): Klein Drakenstein Mountains near Salem, 
Galpin 10594. 3321 (Ladismith): between Calitzdorp and 
Cango Caves, Hutchinson 1145. 3322 (Oudtshoom): 5 km 
E. of George, Hutchinson 1261. 3323 (Knysna): Groot 

River, Fourcade 1459 (K). 3324 (Steytlerville): Klein 
River, Hankey, Long 1350. 3325 (Port Elizabeth): Enon, 
Uitenhage, Thode A 1091 ; “Zuurbergen by Bontjiesrivier”, 
Drege 6722a. 3326 (Grahamstown): Olifantshoek, Alexan- 
dria, Johnson 1119. 3418 (Simonstown): Somerset West, 
Parker 3734. 

An extremely variable species as regards leaf shape, 
size, texture and degree of pubescence. In forest the species 
may have large, chartaceous leaves with little pubescence, 
while a few hundred metres away in a boulder group in the 
open, the species may have small, coriaceous leaves which 
are densely pubescent. Kiggelaria flavo-velutina Sleumer, 
described from East Africa, probably represents the extreme 
in pubescence of this species: it is doubtfully distinct from 
K. africana. 

5304 5. SCOLOPIA 

Scolopia Schreb. in L., Gen. PI. ed. 8, 1: 335 (1789), nom. conserv.; Sim, For. FI. Cape Col. 125 
(1907); Gilg in Pflanzenfam. ed. 2, 21: 418 (1925); Phill., Gen. ed. 2: 513 (1951); Wild in F.Z. 1: 
276 (1960); Bamps in F.C.B. Flacourt. 39 (1968); Sleumer in Blumea 20: 26 (1972); Wild & 
Vidigal, FI. Mocamb. Flacourt. 17 (1973). Type species: 5. pusilla (Gaertn.) Willd. 

Phoberos Lour., FI. Cochinch. 317 (1790). 

Eriudaphus Nees in Eckl. & Zeyh., Enum. 271 (1835). 

Adenogyrus Klotzsch in App. Ind. Sem. Hort. Berol. 1 (1854). 

Rhamnicastrum Kuntze, Rev. Gen. 1: 45 (1891). 

Armed or unarmed shrubs or trees. Leaves alternate, simple, petiolate or subsessile; blade 
elliptic, ovate or subrotund, entire or toothed; stipules minute, caducous. Flowers bi-sexual, in 
axillary racemes, spikes, fascicles or flowers solitary in axils. Receptacles usually funnel-shaped. 
Sepals 4—6. Petals 4—6, but not always present, alternating with sepals. Disc fleshy, with marginal 
glands. Stamens many, borne on surface of disc, exceeding petals in length at anthesis; anthers 
2-thecous, often apiculate. Ovary superior, 1-locular, with few ovules on 2-5 parietal placentas; 
style simple, longish; stigma subcapitate or 2-5-lobed. Fruit a fleshy berry with several seeds and 
remains of sepals, petals and stamens at base and style at apex. Seeds suborbicular; testa leathery; 
embryo straight; cotyledons flat, elliptic-oblong; endosperm fleshy or somewhat homy. 

About 37 species, natives of Africa, Madagascar, Comores, Mascarenes, Malaysia and Australia; 5 species in Southern 
Africa. The generic name is derived from the Greek for “thorn”. 

Flowers racemose: 

Leaves usually narrowly to broadly obovate, sometimes elliptic or ovate with narrowly cuneate base, apex 
acute, margins entire, repand or distantly crenate-serrate with teeth in upper half, upper surface frequently with 

flaky, waxy covering 1. S. zeyheri 

Leaves elliptic or ovate with cuneate base, apex acuminate, margins closely and regularly callose-serrate or serrulate 

throughout whole length of blade, upper surface without flaky, waxy covering 2. S. mundii 

Flowers solitary or fasciculate: 

Leaves 6-11 mm long; stamens 50 or more 3. S. stolzii 

Leaves 2-4 cm long; stamens 10-25: 

Leaves with 7-13 pairs of veins, apex acute to blunt; flowers pedicellate, 2 or 3 per 


Leaves with 5 or 6 pairs of veins, apex acuminate; flowers sessile, solitary 

.4. S. flanaganii 
.5. 5. oreophila 



1. Scolopia zeyheri (Nees) Harv. in F.C. 
2: 584 (1862), in text; Szyszyl., Polypet. 
Thalam. Rehm. Ill (1887); Warb. in Pflanzen- 
fam. 3,6a: 29 (1893); Dur. & Schinz, Consp. FI. 
Afr. 1: 220 (1898); Sim, For. FI. Cape Col. 126, 
t.2 (1907); Gilg in Bot. Jahrb. 40 : 481 (1908); 
Pflanzenfam. ed. 2,21: 418 (1925); Henkel, 
Woody PI. Natal 73, 90 (1934); Phill. in 
Bothalia 1: 84 (1922); Wild in F.Z. 1: 276 
(1960); l.c. 565 (1961); Bamps in F.C.B. 
Flacourt. 40 (1968); Exell in F.Z. 3: 141 (1970); 
Sleumer in Blumea 20: 60 (1972); Wild & 
Vidigal in FI. Mocamb. Flacourt. 17 (1973). 
Type: Cape, Olifantshoek and Addo, Ecklon & 
Zeyher 1756 (B, holo.t; P!; L; SAM!). 

Eriudaphus zeyheri Nees in Eckl. and Zeyh., Enum. 2: 
272 (1836). E. ecklonii Nees, l.c. 271 (1836). Type: Cape, 
Kat River, Ecklon & Zeyher 1754 (B, holo.t; G! ; K!; P!; 

Phoberos zeyheri (Nees) Am. in Hook., J. Bot. 3: 150 
(1841); Presl, Bot. Bemerk. 70 (1844); Pappe, Silv. Cap. 4 
(1854); Harv. in F.C. 1: 68 (1860). P. ecklonii (Nees) Am. 
ex Presl, Bot. Bemerk. 70 (1844); Pappe, Silv. Cap. 3 
(1854); Harv. in F.C. 1: 68 (1860). 

Scolopia gerrardii Harv. in F.C. 2: 584 (1862); Dur. 
and Schinz, Consp. FI. Afr. 1: 220 (1898): Gilg in Bot. 
Jahrb. 40: 481 (1908); Engl., Pflanzenw. Afr. 3, 2: 576 
(1921); Gilg in Pflanzenfam. ed. 2, 21: 418 (1925). Type: 
Natal, Nototi River, Gerrard & McKen 26 (TCD, holo. !; 
PRE, photo.). S. ecklonii (Nees) Harv. in F.C. 2: 584 
(1862), in text; Szyszyl., Polypet. Thalam. Rehm. Ill 
(1887); Warb. in Pflanzenfam. 3, 6a: 29 (1893), as S. 
ecklonii (Am.) Warb.; Dur. and Schinz, Consp. FI. Afr. 1: 
220 (1898); Gilg in Bot. Jahrb. 40: 481 (1908); Phill. in 
Bothalia 1: 86 (1922); Gilg in Pflanzenfam. ed. 2, 22: 420 
(1925); Burtt Davy, FI. Transv. 1: 215 (1926); Henkel, 
Woody PI. Natal 58, 73, 80 (1934). S. engleri Gilg in Bot. 
Jahrb. 40: 481 (1908); Engl., Pflanzenw. Afr. 3, 2: 577 
(1921); Gilg in Pflanzenfam. ed. 2,21: 420 (1925). Type: 
Transvaal, Wonderfontein, Engler 2882a (Bt). S. ecklonii 
(Nees) Harv. var. gerrardii (Harv.) Phill. in Bothalia 1: 86 
(1921). — var. engleri (Gilg) Phill., l.c. 86 (1921); Burtt 
Davy, FI. Transv. 1: 215 (1926). S. thorncroftii Phill., l.c. 
84 (1921); Burtt Davy, l.c. 215 (1926.) Type: Transvaal, 
Barberton, Thorncroft 811 (NH, holo.!; BM!). 

Rhamnicastrum zeyheri (Nees) Kuntze, Rev. Gen. 1: 
45 (1891). R. ecklonii (Nees) Kuntze, l.c. 45. 

Shrub or tree up to 13 m high, unarmed or 
with straight branched or unbranched rather 
stout spines up to 15 cm long. Bark dark grey, 
fissured. Leaf-blade narrowly to broadly ob- 
ovate, sometimes elliptic or broadly elliptic to 
ovate, 3,5-10 cm long, 1,2-6 cm wide, apex 
acute, obtuse or emarginate, base narrowly 
cuneate to cuneate, margin entire, repand or 
distantly crenate-serrate, teeth usually in upper 
half only, coriaceous, dull green above, paler 
below, often bright pink when young, upper 
surface frequently covered with a thin flaky, 

waxy covering, veins 3-5 pairs, sometimes 
appearing 3-nerved at base, somewhat project- 
ing, glabrous; petioles up to 1,5 cm long. 
Flowers in axillary racemes, 2,5-5 cm long; 
pedicels 3-5 mm long with small triangular 
bracts at base. Receptacle broadly funnel- 
shaped. Sepals 5-6, elliptic, 1,3-1, 5 mm long, 
0,75-1 mm wide, acute to round, margin 
sparsely ciliate. Petals (when present) narrowly 
elliptic, 1,2 mm long, 0,5-0,75 mm wide, 
acute. Disc annular, with small fleshy lobes at 
margin, densely villous. Stamens numerous; 
filaments 2-2,5 mm long; anthers oblong, 0,75 
mm long, arcuate, apiculate. Ovary ovoid to 
subglobose, 1,3 mm diam., sulcate, glabrous; 
ovules 4; style 2 mm long; stigma bifid, 
subsessile. Fruit globose, up to 9 mm in diam., 
glabrous, with persistent style; seeds 4, sub- 
ovoid, 4 mm long, 3 mm wide, angular, 
minutely reticulate. 

A tree found in bush clumps, scrub and forest from the 
coast to the upland regions of the Transvaal, Orange Free 
State, Swaziland, Natal and the Cape. Also widespread in 
tropical Africa occurring as far north as Cameroon, Uganda 
and Kenya. 

Transvaal. — 2329 (Petersburg): Houtbosch, Rehmann 
6464 (K; BM). 2330 (Tzaneen): Westfalia Estates, 
Duiwelskloof, Scheepers 1069. 2428 (Nylstroom): Mos- 
dene, Naboomspruit, Galpin M691 . 2430 (Pilgrims Rest): 
The Downs, Renny DB23. 2526 (Zeerust): Wonderfontein, 
Burn l)a\\' 7564 (K). 2527 (Rustenburg): ‘‘Rainhill” near 
Rustenburg, Codd 1083. 2529 (Witbank): Maleeuw Kop, 
Acocks 23351. 2531 (Komatipoort): Klokwene, Kruger 
National Park, Van der Schijff & Marais 3784. 2628 
(Johannesburg): Prosser 1551 (K). 2630 (Carolina): Mooihoek, 
Devenish 881 . 

O.F.S. — 2627 (Potchefstroom): 8 km W. of Parys, 
Stapleton s.n. 2827 (Senekal): Willem Pretorius Game 
Reserve, Muller 895. 2828 (Bethlehem): Farm Suzanna, 
Scheepers 1788. 

Swaziland. — 2631 (Mbabane): near Hlatikulu, Comp- 
ton 31337; Mafuteni Hill, Manzini, Compton 31664; 
Tungulu Mountain, Mankaiana, Miller S/266. 

Natal. — 2632 (Bela Vista): Manguzi Forest W. of 
Maputa, Tinley 338. 2729 (Volksrust): Farm Glendale, 
Normandien, Biggs 51. 2731 (Louwsburg): Ngome Forest, 
Gerstner 4881. 2732 (Ubombo): False Bay Park, Ward 
7059. 2830 (Dundee): 19 km from Muden, Moll 3242. 2831 
(Nkandla): Ngoye Forest, Wells & Edwards 66. 2832 
(Mtubatuba): Hluhluwe Game Reserve, Ward 2635. 2929 
(Underberg): Estcourt, above New Formosa, A cocks 10676. 
2930 (Pietermaritzburg): Westville, Wood 8597 (K). 2931 
(Stanger): Groutville, Moll 2531. 3030 (Port Shepstone): 
Ifafa River Lagoon, Moll 3587. 

Cape. — 3028 (Matatiele): Pot River Berg, Galpin 6830. 
3029 (Kokstad): Mt. Ingeli, Nicholson 759. 3126 

(Queenstown): Fincham’s Nek, Galpin 1822. 3129 (Port St. 
Johns): lntafufu, Cooper 3129. 3130 (Port Edward): 
Nyameni Mouth, Strey 8617. 3225 (Somerset East): 
Bruintjies Hoogte, Burchell 3069 (K). 3226 (Fort 

Beaufort): Kat River, Ecklon & Zeyher 1754 (K). 3227 



(Stutterheim): Keiskammahoek, Killick 894. 3228 (Butter- 
worth): Kentani, Pegler 2109. 3322 (Oudtshoom): Wilder- 
ness, Compton 10710 (NBG). 3323 (Willowmore): hills S. 
of De Vlugt, Fourcade 3774 (K). 3325 (Port Elizabeth): 1,6 
km N. of Zuurberg Inn, Johnson 738 (K). 3326 

(Grahamstown): near Port Alfred, Schlechter 2733 (K; BM; 
P; G). 3327 (Peddie): Green Point, East London, Smith 
3779. 3422 (Mossel Bay): Mossel Bay, Prior s.n. 3423 
(Knysna): Buffalo Bay, Knysna, Taylor 442 (NBG). 

A variable species as regards growth form and 
leaf-shape. This variability accounts for the extensive 
synonymy of the species, especially for tropical Africa. See 
Sleumer’s monograph for full synonymy. There has been 
considerable inconsistency in the citation of authors for S. 
zeyheri. The correct citation is (Nees) Harv., but the 
following have been frequently used: (Am.) Warb., (Am.) 
Harv., (Nees) Szyszyl. S. zeyheri is known by several 
common names, the best-known probably being Thom 
Pear. The wood is very hard and was once used in 
wagon-making chiefly for axles, felloes and spokes. The 
extreme hardness of the wood also rendered it useful for the 
manufacture of teeth for mill wheels. 

2. Scolopia mundii (Eckl. &Zeyh.) Warb. 
in Pflanzenfam. 3,6a: 29 (1893), as (Am.) 
Warb.; Dur. & Schinz, Consp. FI. Afr. 1: 220 
(1898); Sim, For. FI. Cape Col. 26, t.l. (1907); 
Gilg in Bot. Jahrb. 40: 481 (1908); Engl., 
Pflanzenw. Afr. 3,2: 576 (1921); Phill. in 
Bothalia 1: 85 (1922); Gilg in Pflanzenfam. ed. 
2,21: 418 (1925); Burtt Davy, FI. Transv. 1: 
215 (1926); Henkel, Woody PI. Natal 73, 77 
(1934); Sleumer in Blumea 20: 59 (1972); 
Killick in Bothalia 11: 280 (1974), Type: Cape, 
“Sitsikamma”, Mund s.n. (B, holo.f; K!; S!). 

Eriudaphus mundii Eckl. & Zeyh., Enum. 2: 272 
(1836); Steud., Norn. Bot. ed. 2: 589 (1840); Walp., Repert. 
1: 548 (1842). E. serratus Harv., Gen. PI. 1: 417 (1838); 
Dur. & Schinz, Consp. FI. Afr. 1: 220 (1898). Types: Cape, 
Van Stadensberg, apparently syntypes ar e Zeyher 788 (K!; 
BM!; TCD); Zeyher 3785 (LD!; P!; S!; SAM!; TCD; Z!). 

Phoberos mundii (Eckl. & Zeyh. ) Presl, Bot. Bemerk. 70 
(1844); Pappe, Silv. Cap. 3 (1854); Harv. in F.C. 1: 68 

Adenogyrus krebsii Klotzsch, App. Ind. Sem. Hort. 
Berol. 1 (1854); Walp., Ann. 4: 227 (1857). Type: “In 
promontorio bonae spei”, Krebs s.n. (Bf). 

Rhamnicastrum mundii (Eckl. & Zeyh.) Kuntze, Rev. 
Gen. 1: 45 (1891). 

Shrub or tree up to 16 m high (27 m fide 
Sim, l.c. ), unarmed or with simple, slender 
spines up to 4 cm long. Bark thin, grey or light 
brown, smooth when young, thick, grey, brown 
or yellowish, rough, fissured when old. 
Branches sometimes with axillary galls re- 
sembling fruits. Leaf-blade elliptic, ovate or 
rarely subrotund, acuminate at apex, cuneate to 
rounded at base, (2,5) 4-7 cm long, 2-4 cm 
wide, margin closely and regularly callose ser- 
rate or serrulate, sometimes with spinose teeth. 

midrib channelled above, prominent below, 
veins 4-6, Vi -angled, looping some distance 
from margin, subprominent, dark green and 
shiny above, paler below, coriaceous, glabrous; 
petiole 3-10 mm long, reddish. Flowers 
greenish white, in racemes up to 3 cm long, 
pedicels up to 6 mm long. Sepals 4 or 5, ovate 
to broadly ovate, apex acute or obtuse, 2-2,2 
mm long, 1,5-2 mm wide. Petals when present, 
deltoid, 1,2 mm long, 1 mm wide. Disc annu- 
lar, with small, orange, fleshy lobes, 0,75 mm 
long, densely villous. Stamens numerous, fila- 
ments 4—5,5 mm long, glabrous; anthers ob- 
long, 0,7 mm long, arcuate, apiculate. Ovary 
ovoid, 1,5 mm long, 1,5 mm wide at base, 
sulcate, glabrous; ovules several; style 3,5 mm 
long, sulcate; stigma bilobed with each lobe 
again faintly bilobed. Fruit globose, 7-10 mm 
diam., glabrous, with persistent style, red; seeds 
3 or 4, angular, 4 mm long, 2 mm wide. Fig. 19. 

A forest tree occurring in the Transvaal, Orange Free 
State, Swaziland, Natal, Lesotho and the Cape as far south 
as the Cape Peninsula. 

Transvaal. — 2430 (Pilgrims Rest): Mt. Sheba Forest, 
Jones 67. 2527 (Rustenburg): Tonguane Kloof, De Winter 
s.n. 2730 (Vryheid): Mooihoek, Devenish 1024. 

O.F.S. — 2828 (Bethlehem): farm of Mr Naude, Theron 
2183. 2829 (Harrismith): Maweni Heights, Van Zinderen 
Bakker Jnr. 12. 

Swaziland. — 2631 (Mbabane): 5 km E. of Mbabane, 
Miller S/215. 

Natal. — 2730 (Vryheid): Donkerhoek, Devenish 1311. 
2828 (Bethlehem): Royal Natal National Park, Hutchinson 
4497 (K); Galpin 9514. 2829 (Harrismith): Cathedral Peak 
Forest Station, Killick 1739. 2830 (Dundee): Qudeni, Davis 
89 (NH). 2832 (Mtubatuba): Enseler.i Nature Reserve, 
Venter 594. 2929 (Underberg): Ntabamhlope, Pentz 312. 
2930 (Pietermaritzburg): Howick, Moll 1070; Dargle, Moll 
864. 2931 (Stanger): Hawaan Forest, Ross & Moll 2296. 
3030 (Port Shepstone): Valleys Nature Reserve, Nicholson 

Lesotho. — 2927 (Maseru): Mamathes, Marais 1084; 
Jacot Guillarmod 469; bank of Mohale River, Jacottet sub 
Dieterlen 1062; Matukeng, Dieterlen 852. 

Cape. — 3028 (Matatiele): 5,3 km S. of Mt. Frere, Story 
942. 3029 (Kokstad): 6,9 km N. of Emagusheni Store, 
Marais 1164. 3126 (Queenstown): Hangklip Mountain, 
Roberts 2121. 3128 (Umtata): Baziya Forest, Marais 497. 
3226 (Fort Beaufort): E. of Abies Grove, Hogsback, Wells 
3779. 3227 (Stutterheim): Mt. Kemp, Killick 908. 3228 
(Butterworth): near Willowvale, A cocks 12278. 3318 (Cape 
Town): Kirstenbosch, Esterhuysen 18637. 3323 (Willow- 
more): Mund s.n. (K). 3325 (Port Elizabeth): Van Stadens 
Mountain, Zeyher 788 (BM; K); Zuurberg Sanatorium, 
Long 1251. 3326 (Grahamstown): Grahamstown, Mac- 
Owan 957 (BM; K). 3327 (East London): Fort Grey, Forest 
Reserve, Wells 3917. 3423 (Knysna): Knysna, Keet 512. 

Known as the Red Pear (Rooipeer) or Mountain 
Safraan (Berg Safraan) The wood is hard, heavy and close 
grained and was formerly used in wagon making, chiefly for 
felloes. It is recommended by Galpin (fide Burtt Davy l.c.) 
as a hedge plant. 



r- — 'ifeSSSTT 

Fig. 19. — 1, Scolopia mundii, flowering twig, (Ranger s.n.); a, fruiting twig, x 1 (Johnson 979); b, flower, x 4 (Ranger 
s.n.); c, portion of flower with sepals excised to show annular disc, x 5 (Ranger s.n.); d, anther, x 20 (Ranger s.n.). 



3. Scolopia stolzii Gilg in Engl., Pflan- 
zenw. Afr. 3,2: 577 (1921) Pflanzemfam. ed. 2,21: 
420 (1951); Sleumer in Notizbl. Bot. Gart. Berl. 
12: 142 (1936); Brenan, Checklist Tang. Terr. 
236 (1949); Wild in F.Z. 1: 278, t.47A (1960); 
F. White, For. FI. N. Rhod. 267 (1962); Bamps 
in F.C.B. Flacourt. 39 (1968); Sleumer in 
Blumea 20: 51 (1972); Wild & Vidigal in FI. 
Mocamb. Flacourt. 18 (1973); Killick in 
Bothalia 11: 280 (1974). Type: Tanzania, 
Kyimbila, Kiwira River, Stolz 1742 (B, holo.f; 
BM; K; PRE!). 

S. riparia Mildbr. & Sleumer in Notizbl. Bot. Gart. Berl. 
1 1: 1077 (1934); Brenan, Checklist Tang. Terr. 235 (1949). 
Type: Tanzania, Tandala Kinga Mts., Iletile River, Stolz 
2200 (B, holo.-j-; L; M!). S. stolzii Gilg var. riparia (Mildbr. 
& Sleumer) Sleumer in Blumea 20: 52 (1972). 

Tree up to 14 m high, 25 cm d.b.h., 
unarmed, trunk brownish with irregularly 
shaped exfoliation sloughing to leave dark 
brown shallow basins; twigs white to pale grey, 
lenticellate. Leaves exstipulate, elliptic or 
ovate, 6-11 cm long, 3,4— 5,7 cm wide, apex 
obliquely acuminate to obtuse, base broadly 
cuneate, margin remotely callose subserrate to 
crenulate, 5-9 pairs of nerves, basal 2 pairs 
acutely angled (curved ascending), remainder 
obtusely angled, looping before margin, im- 
mersed above, projecting below, coriaceous, 
glabrous, darkish green above, paler below, 5-8 
mm long, recurved. Flowers solitary or in pairs, 
sessile, in upper leaf axils. Bracts 5, deltoid, 
0,5 mm long, somewhat cartilaginous. Sepals 
5, approximately triangular, 1,4—2 mm long, 
ciliate. Petals 5, resembling sepals, but some- 
times rounded, early caducous. Disc annular, 
regularly lobed, 0,5 mm long, “salmon pink”. 
Stamens numerous, sometimes over 100; fila- 
ments 7,5 mm long; anthers 0,75 mm long, 
slightly apiculate. Ovary ovoid, 3,5 mm long 
with longish white hairs; placentae 5-6 with 
numerous ovules; style short, 5 or 6 branched, 
branches 1,8 mm long, channelled above, 
glabrous. Fruit subglobose, 2,5 cm diam., 
fleshy, often crowned with persistent style; 
seeds 8-17, oblong, c. 5 mm long, brown. Fig. 
20 . 

Fairly widespread in tropical Africa occurring as far 
north as Cameroon. In Southern Africa known only from 
the Sihadla River Crossing and Mtunzini in Natal, where it 
grows in swamp and dune forest respectively. 

Natal. — 2732 (Ubombo): Sihadla River Crossing, Kil- 
lick & Vahrmeijer 4061 ; Moll <6 Nel 5608; Moll 5724; 
Strey8187; Strey & Moll 3890. 2831 (Nkandla): Mtunzini, 
Umlalazi Lagoon, Venter 6013. 

Only recently discovered in Southern Africa. Accord- 
ing to a note on Moll & Nel 5608 the foliage is pendent 
with the young leaves dark pink and highly conspicuous and 
the slash is pale pink. A variable species as regards leaf size, 
texture, margin and pubescence of ovary. 

S. stolzii Gilg is treated as a nomen nudum by Wild & 
Vidigal (1973), but the present author has followed Sleumer 
(1972) in accepting the 3-line description as adequate for 
valid publication. 

4. Scolopia flanaganii (H. Bol.) Sim, 
For. FI. Cape Col. 127 t.3 (1907); Gilg in Bot. 
Jahrb. 40: 484; Engl., Pflanzenw. Afr. 3,2: 576 
(1921); Phill. in Bothalia 1: 84 (1921); Gilg in 
Pflanzenfam. ed. 2, 21: 418 (1925); Henkel, 
Woody PI. Natal 80 (1934); Sleumer in Blumea 
20: 50 (1972). Type: Cape, near Komga, 
Flanagan 682 (BOL, holo.!; K!; NH!; PRE!; 

Xylosma flanaganii H. Bol. in J. Bot. Lond. 34: 17 

Tree or shrub up to 5 m high with trunk up 
to 17 cm d.b.h., armed with slender spines up to 
3 cm long chiefly on coppice shoots and young 
growth. Branchlets glabrous or puberulous. 
Leaf -blade: adult leaves narrowly elliptic to 
elliptic or narrowly obovate to obovate, 2-3,5 
cm long, 0, 6-2,2 cm wide, apex acute to 
obtuse, base narrowly to broadly cuneate, mar- 
gins serrulate or remotely serrulate, often entire 
in lower l h, teeth callose, coriaceous, glabrous, 
midrib prominent on lower surface, lateral 
nerves 7—13, subimmersed; petiole 2-4 cm 
long; juvenile leaves ovate, 1, 8-3,4 cm long, 
1, 1-2,2 cm wide, apex acute, base broadly 
cuneate or cordate, margins serrate, sometimes 
shortly spine-tipped, coppice shoot leaves 
broadly elliptic or circular, 2, 6-3, 3 cm long, 
2-2,4 cm wide, apex acute or rounded, base 
broadly cuneate or rounded, margins serrate. 
Flowers solitary or fasciculate in leaf axils, 
5-merous; pedicels 1,25-1,75 mm long, 
stoutish, bracteate. Sepals broadly ovate, 0,5- 
0,75 mm long, 0,75-1 mm wide, concave, 
somewhat keeled dorsally, scariose, ciliate on 
margin. Petals early caducuous, elliptic, ovate 
or circular, 0,75-1 mm long, 0,5-1 mm wide. 
Disc annular of quadrate fleshy lobes, glabrous. 
Stamens 10-20; filaments 2-2,5 mm long; an- 
thers oblong, 0,5 mm long. Ovary ovoid, 1-2,5 
mm long, 1-1,25 mm wide at base, pubescent 
or glabrous; stigma obscurely 2-4-lobed. Fruit 
subglobose, up to 7 mm long, 5 mm wide, 
pubescent, red. Seeds 1 or 2. Fig. 21. 



Fig. 20. — 1, Scolopia stolzii, fruiting twig; a, flower in late bud; b, mature flower; c, young fruit with persistent styles; all x 
4 /s (Killick & Vahrmeijer 4061). 



Fig. 21. — 1, Scolopia flanaganii, series showing variation in leaf shape, all X 1. 1 and 2, coppice shoot leaves (Killick 
4076 and 4068); 3-5, juvenile leaves (Killick 4056, Edwards 4313 and Killick 1957); 6-13, adult leaves (Acocks 
10519, Killick 4056, 4075, 4075, 4070 and 4070, Flanagan 682 and Killick 4069). 



Occurring inside and along the margin of forest and 
scrub from the eastern Cape to the midlands and uplands of 

Natal. — 2929 (Underberg): Giants Castle Game Re- 
serve, Killick 4056; Hlatikulu Forest, Killick 1957; 11 km 
from Bulwer on road to Impendhle, Killick & Marais 2103. 
2830 (Dundee): Qudeni, Gerstner 667: Umhlumba Moun- 
tain, West 1470. 2930 (Pietermaritzburg): Keerom Farm, 
Greytown, Cooper 60 (NH). 

Cape. — 3128 (Umtata): All Saints Nek, Marais 502; 40 
km S. of Ugie, Edwards 4313. 3227 (Stutterheim): near 
Komga, Flanagan 682; Fort Cunynghame, Killick 4075; 
Kei Road, Killick 4069; Mount Kemp, Killick 902. 

S. flanaganii is an extremely variable species espe- 
cially in the shape of the adult, juvenile and coppice shoot 
leaves. Also, the ovaries vary from completely glabrous to 
pubescent. A feature of the leaves is the frequent presence 
of small shot-holes apparently caused by a blue-green alga. 

5. Scolopia oreophila (Sleumer) Killick 
in Bothalia 11: 515 (1975). Type: Natal, Ut- 
recht District, Farm Retirement, Devenish 1319 
(K, holo. !; PRE!; S!). 

S. flanaganii (H. Bol.) Sim var. oreophila Sleumer in 
Blumea 20: 51 (1972). 

Tree up to 13 m high, 23 cm d.b.h., spines 
on young plants and coppice shoots. Branchlets 
glabrous, puberulous at apex. Leaf-blade 

rhombic-elliptic (coppice shoot leaves ovate), 
2-3 cm long, 1-1,7 cm wide, apex acuminate, 
base cuneate (wedge-shaped), margin distantly 
serrate-crenate, teeth callose, midrib prominent 
below, lateral veins 5-6 pairs, subimmersed, 
coriaceous, glabrous; petiole 2—4 mm long, red, 
thickish. Flowers solitary in leaf axils, sessile, 
bracteolate at base, (4) 5-merous. Sepals nar- 
rowly elliptic to broadly ovate, slightly con- 
cave, 2-2,5 mm long, 1-2 mm wide. Petals not 
seen. Disc annular, of orange-coloured, fleshy 
lobes, 0,5 mm long, glabrous. Stamens 15-25; 
filaments 2,5-3 mm long, glabrous; anthers 
oblong, 0,5 mm long, sometimes with short 
black bristles. Ovary ellipsoid to ovoid, 2-3 
mm long, 2 mm wide, glabrous; ovules 3; style 
2,5-3 mm long, glabrous; stigma bilobed with 
each lobe faintly bilobed. Fruit not seen. 

Known only from the Utrecht District of Natal, where 
it occurs along the margin of streambank scrub and forest. 

Natal. — 2730 (Vryheid): Farm Donkerhoek, Devenish 
528; Farm Retirement, Devenish 1319; Killick 4057; Farm 
Naauwhoek, Devenish 1141. 

Can be distinguished from S. flanaganii by the leaves 
which are somewhat rhombic in shape with an acuminate 
apex, 5 or 6 pairs of veins and reddish petioles and the 
sessile, solitary flowers. 


Pseudoscolopia Gilg in Bot. Jahrb. 54: 343 (1917); Phill., Gen. ed. 2: 513 (1951). Type species: 
P. polyantha Gilg. 

Pseudoscolopia Phill., Gen. 416 (1926), nom. illegit. Type species: P. fraseri Phill. 

Shrub or small tree. Leaves opposite, petiolate, exstipulate; blade elliptic, serrate. Inflores- 
cence a lax, axillary cyme. Flowers bisexual. Sepals 4, valvate in bud, persistent. Petals 4, 
subsimilar to sepals, imbricate in bud, persistent. Stamens many; filaments free; anthers arcuate. 
Ovary superior, ovoid-globose, 1 -locular with 2 or 3 parietal placentas and 1 ovule on each 
placenta, pubescent; style terete, 2 or 3-fid at apex. Fruit a 2 or 3-valved capsule, ovoid, with 
persistent style. Seed ellipsoid, covered with stellate hairs; embryo straight; cotyledons flat, 
somewhat obovate; endosperm sparse. 

A monotypic genus recorded from Natal and the Cape. The genus resembles and is closely related to Scolopia, hence 
the name Pseudoscolopia. 

Pseudoscolopia polyantha Gilg in Bot. 
Jahrb. 54: 343 (1917); Phill. & Hutch, in Hook. 
Icon. PI. t. 3 1 19 (1933). Syntypes: Cape, En- 
kweni and Egossa Forest, Bachmann 1712 and 
1713 (Bf); Beyrich 116 and 125 (B|); Mkam- 
bati River, Marais 969 (PRE, neo. !). 

P. fraseri Phill., Gen. 416 (1926). Type: not indicated, 
probably Cape, Transkei, Ntsubane Forest, Fraser sub PRE 
1417 (PRE, holo.!). 

Shrub or tree up to 5 m high. Branchlets 
angled, glabrous. Leaf-blade narrowly elliptic 
to elliptic, 4-6,5 cm long, 2-3 cm wide, apex 
acuminate or acute, base cuneate, margin ser- 
rate with cartilaginous teeth or nearly entire, 
midrib prominent below, lateral veins 5 or 6, 
not very conspicuous, coriaceous, glabrous; 
petiole 3-8 mm long. Inflorescence 4—6- 
flowered, 2-3 cm long. Flowers bisexual; 



Fig. 22.— 1, Pseudoscolopia polyantha, flowering twig, x 1 ( Marais 696); a, flower, x 3 ( Cooper 107); b, gynoecium, x 
8 (Cooper 107); c, longitudinal section through ovary, X 10 ( Cooper 107); d, fruit, x 3 (Esterhuysen 17946); e, seed, 
X 8 (Esterhuysen 17946). 



pedicels 6-11 mm long, puberulous. Sepals 
elliptic, 6-7 mm long, 2,5-3 mm wide, apex 
acuminate, pubescent, margins ciliate. Petals 
subsimilar to sepals. Stamens many; filaments 
3-4 mm long, glabrous; anthers oblong, c. 1 
mm long, prominently arcuate. Ovary ovoid- 
subglobose, 1,5-2 mm long, villous; style ter- 
ete, 3 mm long, glabrous; stigma 2 or 3-fid, 
arms c. 1,5 mm long. Fruit ovoid, 5 mm diam., 
with persistent style. Seeds ellipsoid, 3 mm 
long, stellately pubescent. Fig. 22. 

A forest tree showing wide discontinuity in its distribu- 
tion: one specimen, Esterhuysen 17946, has been collected 

at Rivers Kloof in the Piketberg District of the south- 
western Cape and the rest in Pondoland and southern Natal. 

Natal. — 2831 (Nkandla): Ngoye Forest, Ross 1863. 
2930 (Pietermaritzburg): Laager Farm, Noodsberg, Nichol- 
son 544 (NH). 3030 (Port Shepstone): Beacon Hill, Strey 
6530 ; 7226; Nicholson 817. 3130 (Port Edward): Umtam- 
vuna River George, Cooper 107. 

Cape. — 3129 (Port St. Johns): Mkambati River, Marais 
969; Ntsubane Forest, Fraser sub PRF 3058; 3105; 3133. 
3218 (Clanwilliam): Rivers Kloof, Esterhuysen 17946. 

With the appearance of a Scolopia, but differing in its 
opposite leaves, 2 or 3-ovuled ovary and capsular fruits. 
Pseudoscolopia polyantha bears some resemblance to Cas- 
sipourea flanaganii (Schinz) Alston of the family 
Rhizophoraceae with which it has sometimes been con- 



Gerrardina Oliv. in Hook. Icon. PI. 11: 60, 1. 1075 (1870); Gilg in Bot. Jahrb. 40: 488 (1908); 
Engl., Pflanzenw. Afr. 3,2: 578, t.256a-c (1921); Gilg in Pflanzenfam. ed. 2, 21: 425, t. 191a-c 
(1925); Phill. , Gen. ed. 2: 514 (1951); Wild in F.Z. 1: 287 (1960). Type species: G.foliosa Oliv. 

Scrambling or erect shrubs or trees. Leaves alternate, simple, petiolate, stipulate; blades 
elliptic or ovate, sometimes acuminate, serrulate to serrate or crenato-serrate. Inflorescence a 
few-flowered axillary cyme on a long peduncle. Flowers bisexual. Calyx-tube campanulate; lobes 
5, imbricate, unequal, 2 outer shorter. Petals 5, shorter than calyx and alternating with the 
calyx-lobes, inserted on the margin of the disc, deciduous. Disc cup-shaped, lining calyx-tube. 
Stamens 5, opposite the petals, inserted on margin of disc. Ovary superior, 1 -locular, with 4 
pendulous ovules. Fruit a dry capsule; seeds 1-4, ellipsoid or obovoid, testa smooth or reticulate. 

A genus of 2 species, 1 occurring in Southern Africa and the other in Rhodesia, Malawi, Mozambique and Tanzania. 
The genus was named in honour of W. T. Gerrard, who collected in Natal about the middle of the last century. 

Gerrardina foliosa Oliv. in Hook. Icon. 
PI. 11: 60, 1. 1075 (1870). Gilg in Bot. Jahrb. 40: 
488 (1908); Engl., Pflanzenw. Afr. 3,2: 578, 
t.256a-c (1921); Gilg in Pflanzenfam. ed. 2, 21: 
425, t. 191a-c (1925); Burtt Davy, FI. Transv. 1: 
217 (1926); Henkel, Woody PI. Natal 94 
(1934). Type: Natal, without precise locality, 
Gerrard 1513 (K, holo.!; BM!; PRE, photo.). 

Shrub or tree up to 10 m high. Branches 
greyish-brown to brown, pubescent at ends. 
Leaf-blade elliptic, 2,6-6, 5 cm long, 1-2,5 cm 
wide, apex acute to obtuse, base narrowly to 
broadly cuneate, margin callose-serrulate to 
serrate, sometimes only in upper 2 Is, thickened, 
revolute, midrib prominent below, lateral 
nerves 9-13, subprominent below, tertiary 
nerves reticulate, immersed, glabrescent, 
coriaceous, pale green below; petiole 2-7 mm 
long. Inflorescence an axillary cyme; peduncles 
erect, 1-4 cm long, slender. Flowers 2-6 per 
cyme; pedicels 1-3 mm long. Calyx-lobes yel- 

lowish, unequal, margins entire or glandular- 
denticulate, faintly keeled; 2 outer rounded, c. 
1,5 mm diam.; 3 inner elliptic to obovate- 
spathulate, 2-3 mm long, 2 mm wide. Petals 
elliptic, 2 mm long, 1,5 mm wide, deciduous. 
Stamens 1 mm long with filaments reddish 
distally; anthers c. 0,7 mm long. Disc shallowly 
5-lobed, reddish-brown. Ovary depressed 
ovoid, 8-ridged radially above, 1 -locular; 
ovules 4, pendulous; style 1 mm long, reddish 
distally; stigma capitate, faintly bilobed, free. 
Fruit bright red, a dry capsule with persistent 
style, globose, 5-6 mm diam.; seed 1, brown, 
ellipsoid-obovoid, 4 mm long, 2,2 mm wide, 
testa smooth, brown. Fig. 23. 

Occurs on rocky hillsides, krantzes and in forest in the 
eastern Cape, Natal, Swaziland and the eastern Transvaal. 

Transvaal. — 2430 (Pilgrims Rest): Mamotswiri, Junod 
187; Smuts & Gillett 3573; God’s Window, Mogg 3329. 
253 1 (Komatipoort): Saddleback Range, Barberton, Galpin 
473 (K); Roses Creek, Thorncroft 2066. 



Fig. 23. — 1, Gerrardina foliosa, flowering twig, x 1 (Galpin 3481); a, longitudinal section through fi° wer < ' 
3853); b, longitudinal section through ovary, x 8 (Strey 3853); c, fruit, X 2 A (ex colour slide, Strey). 

6 (Strey 



Swaziland. — 2531 (Komatipoort): King’s Forest, 

Compton 27817; Havelock Mine, Miller 5785. 2631 
(Mbabane): Mahlangatsche Mt., Mankaiana, Miller S/268. 

Natal. — 2831 (Nkandla): Ngoye Forest, Mtunzini, 
Huntley 730. 2930 (Pietermaritzburg): 59 km from 

Wartburg on Tongaat Road, Edwards 3006; Table Moun- 
tain, Killick 146. 3029 (Kokstad): Murchison, Coleman 452 
(NH). 3030 (Port Shepstone): Umtwalumi, Rudatis 584; 
Oribi Gorge, Strey 3853. 3130 (Port Edward): Mtamvuna 
Forest Reserve, Ward 7174. 

Cape. — 3129 (Port St. Johns): Umsikaba River near 
mouth, Marais 1183; top of Eagles Nest, Port St. Johns, 

Schonland 4079. 3130 (Port Edward): Umtentu River Falls, 
Marais 985. 

A distinct species readily distinguishable from its 
tropical African congener, G. eylesiana Milne-Redh., by its 
leaves which are elliptic-acute instead of ovate-acuminate, 
coriaceous instead of chartaceous and the peduncles which 
are considerably stouter. There are two specimens of 
Gerrard 1513 in Herb. Kew. The one with annotations and 
drawings and marked “Type specimen” is clearly the 

5313 8 . HOMALIUM 

Homalium Jacq., Enum. Syst. PL Ins. Carib. 5, 24 (1760); Benth. & Hook.f., Gen. PI. 1,3: 800 
(1867); Sim, For. FI. Cape Col. 220 (1907); Engl., Pflanzenw. Afr. 3, 2: 578 (1921); Gilg in 
Pflanzenfam. ed. 2,21: 425 (1925); Phill., Gen. ed. 2: 514 (1951); Wild in F.Z. 1: 287 (1960); Wild 
& Vidigal, FI. Mocamb. Flacourt. 31 (1973); Sleumer in Bull. Jard. Bot. Belg. 43: 239 (1973). 
Type species: H. racemosum Jacq. 

Trees or shrubs. Leaves alternate (in African species), simple, petiolate; blade ovate, elliptic, 
suborbicular or oblong, sometimes large; stipules absent, minute or large. Inflorescence an axillary 
or terminal raceme or panicle. Flowers bisexual. Calyx-tube turbinate; sepals 7-9, usually narrow, 
persistent. Petals 7-9, inserted in the mouth of the calyx tube, alternating with and similar to the 
sepals but broader, persistent. Stamens solitary or in fascicles of 2-3 or more, alternating with 
glands which are adnate to the base of the sepals, terete; anthers small, didymous. Ovary 
semi-superior, 1-locular with 2-5 parietal placentas; styles 2-5; stigmas simple or capitate. Fruit a 
semi-superior, leathery capsule, 1- or few-seeded, opening at the apex by 2-5 valves. Seeds 
solitary or few. 

A tropical and subtropical genus of about 180 species; 3 species in Southern Africa, two of which, H. dentatum (Harv.) 
Warb. and H. rufescens Benth., belong to subgenus Blackwellia Warb., section Blackwellia Benth., while the third, H. 
abdessammadii Aschers. & Schweinf. belongs to subgenus Homalium , section Homalium. 

According to Wittstein (1856) the name Homalium is derived from the Greek for “similar or equal” in reference to the 
21 stamens, which are arranged in 7 similar or equal bundles. In actual fact, the stamens can be more or less than 21, but 

when arranged in bundles the bundles are usually equal. 

Stamens solitary at base of each petal: 

Leaves broadly elliptic, 5,5-9 cm long, 3, 5-5, 5 cm wide 1. H. dentatum 

Leaves elliptic to broadly elliptic, 1,5-5 cm long, 0,8-3 cm wide 2. H. rufescens 

Stamens in 3’s opposite each petal 3. H. abdessammadii 

1. Homalium dentatum ( Harv .) Warb. in 
Pflanzenfam. 3,6a: 36 (1893); Engl., Pfanzenw. 
Afr. 3,2: 581 (1921); Gilg in Pflanzenfam. ed. 2, 
21: 427 (1925); Wild in F.Z. 1: 289 (1960); 
Sleumer in Bull. Jard. Bot. Belg. 43: 239 (1973); 
Wild & Vidigal, FI. Mocamb. Flacourt. 31 (1973). 
Type: Natal, “near Port Natal”, Gerrard & 
McKen 50 (TCD, holo.!; PRE, photo.). 

Blackwellia dentata Harv. inF.C. 2: 585(1862). Type as for 
H. dentatum. 

Homalium subsuperum Sprague in Kew Bull. 1923: 184 
(1923); Gilg in Pflanzenfam. ed. 2, 21: 427 (1925). Henkel, 

Woody PI. Natal 86 (1934). Type: Cape, Ndindini Forest, 
Transkei, Kaufmann sub PRF 3225 (sub PRE 2571) (K, holo!; 

Tree 6-30 m high; branchlets glabrous or 
tomentulose. Leaf-blade broadly elliptic, 5,5-10 
cm long, 3,5-7 cm wide, apex abruptly acuminate 
often with oblique point, obtuse or rounded, base 
broadly cuneate to rounded, margin crenate- 
serrate, midrib and lateral veins fairly prominent 
below, lateral veins 5-8, Vi-angled, looping near 
margin, coriaceous, glabrous or tomentulose, 
often with tufts of hair in lower axils; petioles up to 



2.2 cm long, glabrous or tomentulose. Inflores- 
cence of axillary and terminal panicles, divari- 
cately branched; peduncles 2,5-4 cm long, 
glabrous or puberulous; pedicels 0,2-2 mm long, 
puberulous. Calyx-tube puberulous; sepals 6-8, 
narrowly elliptic, 1,2-1,75 cm long, 0, 3-0,7 mm 
wide, puberulous to tomentulose; glands adnate to 
base of sepals, round, sessile, brown, tomentulose. 
Petals 6-8, elliptic, 1 ,5-1,8 mm long, 0,8- 1 ,2 mm 
wide, puberulous to tomentulose. Stamens 6-8, 
terete, 0,2-0, 4 mm long. Ovary narrowly conical, 

1.2 mm long, tomentulose; styles 3 or 4, joined at 
base, with free portion 5 mm long and tomentulose 
in lower half; stigmas simple. Fruit a leathery 
capsule, obovoid, 4 mm long, 2,5 mm wide, tardily 
dehiscent, pilose inside; seeds usually one, c. 2 mm 

Found in scrub and forest in the Transvaal, Natal, eastern 
Cape, Rhodesia, Malawi and Swaziland. 

Transvaal. — 2330 (Tzaneen): Piesangskop, 

Duiwelskloof, Scheepers 903. 2430 (Pilgrims Rest): Erasmus 
Pass, Van der Schijff 7305 ; between Weltevreden Asbestos 
Mine and Olifants River, Codd 10566. 2531 (Komatipoort): 
Eureka City, Sheba Hills, Scheepers SKF 4072; near Malelane, 
Codd 8257; Van der Schijff 2507; Ship Mt„ 14 km S.E. of 
Pretorius Kop, Codd & De Winter 5161. Without precise 
locality: Soutpansberg, Worsdell s.n. (K). 

Swaziland. — ? 2632 (Bela Vista): Chilobe, Lebombo 
Mtns .,MHlerS/19; forest bordering Tibilati stream on E. side of 
Lebombo Mtns., Hornby 2835. 

Natal. — 2731 (Louwsburg): Ngome, Gerstner 5185. 2732 
(Ubombo): Ngwalaweni Forest, Gerstner 4017; Dutton & 
Tinley23. 2830(Dundee): Qudeni Forest, Edwards 2648. 2831 
(Nkandla): Nkandla Forest, Edwards 1366. 2832 (Mtubatuba): 
Hluhluwe Game Reserve, Ward 1712 (NH). 2930 (Pieter- 
maritzburg): Greytown, Cooper 61. 2931 (Stanger): Umgeni, 
Hutchinson 1848; Kildare Estates, Damall, Moll 2901 . 3030 
(Port Shepstone): Umbogintwini, Wood 11977 (K). 

Cape.— 3129 (Port St. Johns): Mpande Forest, Miller sub 
PRF 3756; 2,4km N. ofValley View Store, Lusikisiki, /I cocfcv 
13416; near Fort Donald, Sim 2412. 

The leaves of young coppice shoots are much thinner than 
the adult leaves, the petioles are shorter and the hairy pockets in 
the leaf axils are more conspicuous. In the southern part of the 
Kruger National Park there is a form with densely tomentulose 
leaves and branchlets represented by Codd 5257, Codd & De 
Winter 5161, Van der Schijff 2507 , etc. This form resembles H. 
chasei Wild from Rhodesia in its pubescence, but has the 
paniculate inflorescence of H. dentatum. According to a note on 
Hutchinson 1848 the flowers oiH. dentatum “smell like sweet 

2. Homalium rufescens Benth. in J. Linn. 
Soc. 4: 34 (1 July 1859); Sim, For. FI. Cape Col. 
220 t.68, fig. 1 (1907); Wood, Natal Plants t.529 
(1912); Engl., Pflanzenw. Afr. 3,2: 580 (1921); 
Gilg in Pflanzenfam. ed. 2,21: 427 ( 1 925); Flenkel, 
Woody PI. Natal 84 (1934); Sleumer in Bull. Jard. 
Bot. Belg. 43: 259 (1973). Syntypes: Cape, Enon, 

Drege s.n. (K, lecto. !; BM!; L!; S!; PRE, photo.); 
Natal, Port Natal, Gueinzius s.n. (BM!; K!; S!; 
PRE, photo.). 

Pythagorea rufescens E. Mey. ex Am. in Hook., J. Bot. 3: 149 
( 1 840), nomen nudum; Presl, Bot. Bemerk. 42 ( 1 845), nomen 
nudum. P. africana E. Mey. in Drege, Zwei Pfl. Doc. 134, 214 
(1843), nomen nudum. 

Blackwellia rufescens Am. in Hook., J. Bot. 3: 149 
( 1 840), nomen nudum; Presl, Bot. Bemerk. 42 ( 1 845), nomen 
nudum; Harv. in F.C. 1: 72 (May 1860). 

Tree, 2-7 m high. Leaf -blade elliptic to 
broadly elliptic, sometimes ovate or obovate, 
1,5-5 cm long, 0,8-3 cm wide, apex acute to 
rounded, base cuneate to rounded, margin entire 
or irregularly crenate-serrate, wavy, midrib fairly 
prominent above, lateral veins 6, half-angled, 
dark-green and shiny above, paler below, thinly 
coriaceous, glabrous; petiole 3-7 mm long, 
channelled. Inflorescence of axillary, terminal 
panicles, divaricately branched; peduncles 2-3 
mm long, puberulous; pedicels 2-5 mm long, 
puberulous. Calyx-tube pubescent; sepals 8 or 9, 
subulate, slightly shorter than petals, 1 ,4-2,4 mm 
long, 0,25 mm wide, pubescent, margins ciliate; 
glands opposite sepals, club-shaped, 0,4 mm 
long. Petals 8 or 9, elliptic-spathulate, 2-3 mm 
long, 0,75 mm wide, pubescent, margins white- 
ciliate. Stamens 8 or 9, terete, 2,7-3 mm long, 
sparsely hairy in lower half; anthers transversely 
elliptic, 0,4 mm long. Ovary cone-shaped, c. 1,5 
mm long, tomentulose, with few to many ovules; 
styles 4 or 5, free and glabrous in upper half; 
stigmas simple. Fruits pilose inside, few-seeded. 
Fig. 24. 

Occurring in forest and on riverbanks in the eastern Cape 
and Natal. 

Natal. — 2930 (Pietermaritzburg): Inanda, McKen & 
Buchanan 31 (K); Umzinyati Falls, Wood 11463; 1333 (K); 
McLean & Ogilvie sub PRE 28864. 3030 (Port Shepstone): 
Warner Beach, Ward 980; Uvongo, Mogg 13334; 
Josephine Bridge, Umkomaas River, Bayer 771 . 

Cape. — 3227 (Stutterheim): banks of Nahoon River, East 
London, Galpin 5717. 3228 (Butterworth): Qora Bridge, 
Kentani, A cocks 12291 ; Kei Mouth, Schlechter 22167; 
confluence of Ngqageni and Qora Rivers, Willowvale, 
Ward 5771. 3325 (Port Elizabeth): Zuurberg Pass, Ar- 
chibald 5254. 3326 (Grahamstown): Bathurst Research 
Station, Brink 163. 

A clear-cut species with smaller leaves than H. den- 
tatum. Sleumer (l.c.) has selected the Kew specimen of 
Gueinzius s.n. as the lectotype of H. rufescens. He cites the 
specimen as Gueinzius 92, but on none of the Gueinzius 
specimens seen by the author and certainly not the Kew 
specimen did the number 92 appear. 



Fig. 24. — 1, Homalium rufescens, flowering twig, X 1 (Bayer 771); a, flower, X 5 (Ward 980); b, stamen, X 7 (Ward 
980); c, longitudinal section through ovary, x 20 (Ward 980). 



3. Homalium abdessammadii Aschers. 
& Schweinf. in Sber. Ges. Naturf. Freunde 
Berl. 1880: 130 (1880); Gilg in Bot. Jahrb. 40: 
494 (1908); Pflanzenfam. ed. 2, 21: 428 (1925); 
Engl., Pflanzenw. Afr. 3, 2: 581 (1921); Fer- 
nandes & Diniz in Garcia de Orta 5, 2: 252 
(1957); Wild in F.Z. 1: 291 (1960); F. White, 
For. FI. N. Rhod. 265 (1962); Sleumer in Bull. 
Jard. Bot. Nat. Belg. 43: 311 (1973). Type: 
Sudan, “bei Nganye”, Schweinfurth 3954 (B, 
holo.f; K!; L!; P!; PRE, photo.). 

H. macranthum Gilg in Bot. Jahrb. 40: 496 (1908); 
Pflanzenfam. ed. 2, 21: 428 (1925). Type: Mozambique, 
Rovumu River opposite Lissenga Mtn., Busse 1049 (B, 
holo.f; EA). H. wildemanianum Gilg in Bot. Jahrb. 40: 497 
(1908); Pflanzenfam. ed. 2, 21: 428 (1925). Types: Zaire, 
Katanga, Lukafu, Verdick 123 (Bf; BR; lecto. !; PRE, 
photo.); Lofoi, Verdick 130 (Bf; BR!). H. rhodesicum 
Dunkley in Kew Bull. 1934: 182, fig. (1934). Type: 
Zambia, Kafue, Martin 66 (K, holo. !; PRE, photo ). H. 
abdessammadi Aschers. & Schweinf. subsp. wil- 
demanianum (Gilg) Wild in Bol. Soc. Brot. Ser. 2, 32: 57 
(1958); Wild in F.Z. 1: 291 (1960). 

Tree, 4— 10 m high; branchlets glabrous to 
grey-pubescent. Leaf-blade broadly elliptic, 
6-8 cm long, 4—5 cm wide, apex abruptly 
acuminate, acute, obtuse or rounded, base 
broadly cuneate to rounded, margin coarsely 
crenate-serrate; midrib and lateral nerves more 
prominent below, lateral nerves 6-8, ^-angled, 
looping near margin, glabrous except for few 

hairs on midrib, sometimes with hairy tufts or 
pockets in axils of nerves, coriaceous; petiole 
0,8-1, 5 cm long, glabrous or pubescent. Flow- 
ers in lax, terminal panicles up to 17 cm long, 
flowers solitary or 2 together, sessile or subses- 
sile on puberulous branches of panicle. Sepals 
5-7, subulate-triangular, 2,5-3 mm long, 
1,75-2 mm wide, pubescent dorsally, glabrous 
ventrally, margins ciliate. Petals 5-7, elliptic- 
ovate, 3,3 mm long, 2-2,25 mm wide, pubes- 
cent dorsally, glabrous ventrally, margins 
ciliate. Stamens in 3’s opposite petals, 4 mm 
long, glabrous or with long hairs in lower half; 
anthers 0,5 mm long. Disc glands discoid, 1 
mm diam., sessile, tomentulose. Ovary conical, 
pilose outside and inside; style 2-3 mm long 
with diverging branches 0,3 mm long. Fruit 
capsular, woody, surrounded by persistent 
calyx and petals. Seeds usually solitary, nar- 
rowly ovoid, 1 mm long; testa thin, brown. 

Found on riverbanks in tropical Africa, where it is 
fairly widespread, and in the Eastern Caprivi Strip of South 
West Africa. 

S.W.A. — 1724 (Katima Mulilo): Katima Mulilo, West 

Readily distinguished from H. dentatum and H. rufes- 
cens by the stamens, which are in 3’s instead of solitary. A 
gradation in degree of pubescence can be traced from a 
completely glabrous to a densely pubescent condition. For 
this reason, subsp. wildemanianum is here not accepted as a 
distinct taxon. 

5315 9 TRIMERIA 

Trimeria Harv., Gen. PI. 417 (1838); in F.C. 1: 68 (1860); Benth. & Hook.f., Gen. PI. 1,1: 129 
(1862); Harv., Gen. PI. ed. 2: 15 (1868); Sim, For. FI. Cape Co. 132(1907); Engl., Pflanzenw. Afr. 
3,2: 581 (1921); Gilg in Pflanzenfam. ed. 2, 21: 429 (1925); Phill., Gen. ed. 2: 514 (1951); Wild in 
F.Z. 1: 296 (1960); Wild & Vidigal in FI. Mocamb. Flacourt. 38 (1973). Type species: T. trinervis 

Monospora Hochst. in Flora 24: 660 (1841). 

Renardia Turcz. in Bull. Soc. Nat. Moscou 31, 1: 466 (1858). 

Shrubs or trees. Leaves alternate, simple, petiolate; blade ovate or orbicular, 3-9-nerved from 
base; stipules sometimes foliose, caducous. Flowers dioecious, small, in axillary racemes, spikes 
or panicles. Male flowers: Sepals 3-5, with marginal disc glands opposite each sepal. Stamens 
9-16, inserted in 3’s or 4’s alternating with disc glands; anthers small, subglobose. Female 
flowers: Sepals and petals more or less as in male. Ovary superior, 1 -locular, with 1 or 2 ovules on 
3 parietal placentas; styles 3, short, persistent. Fruit a dry, 3-valved capsule, 1-3-seeded. Seed 
ellipsoid, testa leathery. 



A genus of five species occurring in southern Africa, Rhodesia and tropical east Africa. The generic name is derived 
from the Greek meaning 3 parts in reference to the trimerous sepals and petals of the type species, T. trinervis. 

Sepals and petals trimerous; leaves elliptic or obovate, 2,5-5 cm long, 1,5-2, 5 cm wide 1 . T. trinervis 

Sepals and petals 4 or 5-merous; leaves circular or broadly obovate, 4—11 cm long, 4—10 cm 

wide 2. T. grandifolia 

1. Trimeria trinervis Harv., Gen. PI. 
417 (1838); in F.C. 1: 69 (1860); Sim, For. FI. 
Cape Col. 133, t.8, II (1907); Engl., Pflanzenw. 
Afr. 3, 2; 582 (1921); Gilg in Pflanzenfam. ed. 
2, 21: 430 (1925); Henkel, Woody PI. Natal 107 
(1934). Type: Cape, Van Stadens Mountains, 
Zeyher 724 (TCD, holo. !; PRE, photo.; BM!; 
K!; LD!; SAM!). 

Tree up to 7 m high. Branchlets glabrous 
or pubescent. Leaf-blade elliptic or obovate, 
2,5-5 cm long, 1, 5-2,5 cm wide, apex acumi- 
nate, acute or rounded, base broadly cuneate to 
rounded, margin serrate-callose, 3-(5)-nerved 
at base, nerves conspicuous on lower surface, 
glabrous, subcoriaceous; petiole 0,4—1 cm long, 
glabrous or pubescent. Inflorescence of simple 
spikes, up to 2 cm long; bracts 3, broadly 
deltoid, 0,4 mm long, 6 mm wide, red; rhachis 
glabrous or puberulous. Male flowers sessile. 
Sepals 3, concave, oblong, 0,6 mm long, 0,7 
mm wide, margin sparsely ciliate. Petals 3, 
concave, obovate-spathulate, 1,2 mm long, 1 
mm wide, margin ciliate. Stamens 9, in 3’s at 
base of petals, 1-2 mm long, glabrous; anthers 
small, subglobose. Style aborted, terete, 0,8-10 
mm long. Disc glands opposite sepals, tooth- 
like, fleshy. Female flowers sessile. Sepals 
subrotund, 0,5 mm long, 0,6 mm wide, margin 
sparsely ciliate. Petals 3, subrotund, 0,5 mm 
diam., margin ciliate. Disc glands as in male 
flowers. Ovary ellipsoid, glabrous; styles 3, 0,3 
mm long, slightly diverging, glabrous. Fruits 
obovoid, 4 mm long, 3,8 mm wide; seeds 1, 
ellipsoid-ovoid, 2,5 mm long, 1,3 mm wide; 
testa minutely tesselated. 

A small tree occurring in forest and scrub-forest in the 
Transvaal, Natal and the Cape. There is a 400 km gap in the 
distribution of this species between Cape Town and Knys- 
na. The fact that apparently only one specimen, De Castel- 
nau 395, has ever been collected in the region of Cape 
Town, a well-collected area, makes this record somewhat 
suspect, especially when it is known that De Castelnau 
travelled as far east as “British Kaffraria”. 

Transvaal. — 2630 (Carolina): Mavieristad, Pott 5115. 

Natal. — 2829 (Harrismith): Farm Glendale, Norman- 
dien, Biggs 25. 2929 (Underberg): above Dalton Bridge, 
Wright, West & Acocks 11 (NH). 2930 (Pietermaritzburg): 
8 km along Merrievale/Boston Road, Moll 935. 

Cape. — 3028 (Matatiele): Pot River Berg, Galpin 6575. 
3125 (Queenstown): Gwategu, Galpin 8254. 3225 (Somer- 

set East): Waterkloof Valley, Van der Walt 116. 3227 
(Stutterheim): Mount Kemp, Killick 907. 3318 (Cape 
Town): “circa Cape Town”, De Castelnau 395 (P). 3323 
(Knysna): Knysna, Bowie s.n. (K). 3323 (Port Elizabeth): 
Van Stadens Mt., Zeyher 724 (K); 3989. 3326 

(Grahamstown): Gameston, c. 20 km from Grahamstown, 
Story 2626. 

Easily distinguished from T. grandifolia by the trimer- 
ous flowers and smaller, 3-(5) digitately nerved, elliptic 

2. Trimeria grandifolia (Hochst.) Warb. 
in Pflanzenfam. 3, 6a: 37, f. 13 H-J (1893); Dur. 
& Schinz, Consp. FI. Afr. 1: 225 (1898); Burtt 
Davy, FI. Transv. 1: 219 (1926); Henkel, 
Woody PI. Natal 115 (1934); Killick in Bothalia 
10: 568 (1972); Wild & Vidigal in FI. Mocamb. 
Flacourt. 38 (1973). Type: Natal, “Port Natal”, 
Krauss s.n. (B, holo.f; M, iso.!). 

Monospora grandifolia Hochst. in Flora 24: 661 (1841). 
Type as above. M. rotundifolia Hochst., l.c. Type: Cape, 
Outeniqua and Goukamma River, Krauss s.n. 

Antidesma alnifolium Hook., Icon. PI. t.48 1 (1842). 
Syntypes: Eastern Cape, Bowie s.n. (K!; PRE, photo.); 
Natal, “Port Natal”, Krauss 160 (G!; K!; PRE, photo.). 

Renardia lejocarpa Turcz. in Bull. Soc. Nat. Mosc. 31, 1: 
466 (1858). Type: without precise locality, Ecklon s.n. 

Trimeria alnifolia (Hook.) Harv. in F.C. 1: 69 (1860); 
Sim, For. FI. Cape Col. 132 (1907). T. rotundifolia 
(Hochst.) Gilg in Pflanzenw. Afr. 3, 2: 582 (1921); Milne- 
Redh. in Kew Bull. 1939: 34 (1939); Wild in F.Z. 1: 296 

Tree, 3-10 m high. Bark light brown, 
smooth or longitudinally fissured. Branchlets 
glabrous or pubescent. Leaf -blade circular or 
broadly obovate, 4— 11 cm wide, apex rounded, 
emarginate, bilobed or apiculate, base truncate 
or shallowly cordate, margin serrate, digitately 
5-7-nerved, nerves prominent below, glabres- 
cent with hairs often persisting on nerves, 
subcoriaceous; petiole 1-3 cm long, glabrous or 
pubescent; stipules foliaceous, reniform, 1,2-2 
cm long, 1-2,5 cm wide, caducous, pubescent. 
Inflorescence axillary, branches densely pubes- 
cent; pedicels up to 2,5 mm long, pubescent, 
jointed; bracts deltoid, 1 mm long. Male 
inflorescences in panicles, up to 9 cm long; 
female inflorescences spicate, but sometimes 
branched, up to 8 cm long. Male flowers: 
Sepals 4 or 5, concave, subulate, 1-1,2 mm 
long, 0,5 mm wide, pubescent. Petals 4 or 5, 
concave, obovate-deltoid, 1, 1-1,3 mm long, 



:al T 

Fig. 25. — 1, Trimeria grandifolia, twig with 9 flowers, x 1 (Guy 73); a, longitudinal section through 9 flower, X 5 (Wells 
2136); b, fruit, x 8 (Guy 73); c, seed, X 8 (Guy 73); d, portion of male inflorescence, X 1 (Harrison 258); e, male 
flower, x 10 (Harrison 258). 



0,6-0, 8 mm wide, pubescent. Disc of marginal, 
fleshy, undulate glands opposite sepals. Sta- 
mens 9-16, perigynous, inserted in 3’s or 4’s 
alternating with glands; filaments 2,5 mm long, 
glabrous or pubescent; anthers small, sub- 
globose; style aborted, terete, glabrous. Female 
flowers: Sepals 4, concave-deltoid, 1 mm long, 
0,8 mm wide, pubescent. Petals as sepals. Disc 
of marginal scale-like glands. Ovary ellipsoid- 
obovoid, glabrous; styles 3, 0,5 mm long, 
slightly divergent. Fruits obovoid, 5 mm long, 
3 mm wide, 3-valved; seeds 1 or 2, ellipsoid, 
1,5-2 mm long, 1-1,4 mm wide, testa minutely 
tesselated, with axil on one side. Fig. 25. 

A small tree found in forest from Knysna in the Cape 
eastwards to Natal and northwards to the Transvaal, Swazi- 
land and Rhodesia. 

Transvaal. — 2230 (Messina): Entabeni, Obermeyer 
862. 2329 (Pietersburg): Lejuma near Louis Trichardt, De 
Winter 6016. 2330 (Tzaneen): Duiwelskloof, Scheepers 
552. 2430 (Pilgrims Rest): Mariepskop, Van der Schijff 
4968. 2530 (Lydenburg): Mount Anderson, Galpin 13625. 
2531 (Komatipoort): camp between Louw’s Creek and 
Maid of Mist Mountain, Hutchinson 2444. 

O.F.S. — Without precise locality. Cooper 1008. 

Swaziland. — 2531 (Komatipoort): Piggs Peak Forest, 
Compton 28220. 2631 (Mbabane): Hlatikulu, Compton 

Natal. — 2632 (Bela Vista): Sihadla River, Strey & 
Moll 3903. 2731 (Louwsburg): Ngotshe District, Gerstner 
5181. 2829 (Harrismith): Mazonjwana River, Cathedral 
Peak, Killick 1863. 2830 (Dundee): Umhlumba Mt., West 
1479 (NH). 2831 (Nkandla): Ngoye Forest road, Mtunzini, 
Wells & Edwards 16. 2832 (Mtubatuba): Dukuduku East, 
Moll 2708. 2929 (Underberg): Tabamhlope, West 1140. 
2930 (Pietermaritzburg): Byrne, Galpin 11937. 2931 
(Stanger): 3 km from Mandini on Tugela Mouth road, 
Edwards 1612. 3030 (Port Shepstone): Ifafa, Rudatis 498. 

Cape. — 3028 (Matatiele): Pot River Berg, Galpin 6576 
(K). 3029 (Kokstad): Insizwa Forest, Strey 10728. 3127 
(Lady Frere): Bloemvlei, Elliot, Van Zinderen Bakker 62. 
3129 (Port St. Johns): Lusikisiki District, Galpin 9354 (K). 
3226 (Fort Beaufort): Menziesberg, Scully 599 (K). 3227 
(Stutterheim): Kei Road, Comins 1409. 3228 (Butterworth): 
Kentani, Pegler 746. 3322 (Oudtshoom): Wilderness 
woods, Whellan 1511. 3323 (Willowmore): Storms River 
Forest Reserve, Dahlstrand 479; Deepwalls Forest Re- 
serve, Bos 854. 3327 (Peddie): Amalinda, Ndnni 95. 

Characterized by the large, usually circular, digitately 
5-7-nerved leaves and 4-merous petals and sepals. Accord- 
ing to a note on West 1 140 the plant can behave as a liane. 
T. grandifolia is known to the Europeans as the Wild 
Mulberry or Wilde Moerbei and to the Zulus as iNde- 
blehlovu (ears of an elephant). According to Sim (l.c. 132), 
the tree is usually evergreen, but occasionally deciduous. 
He also states that the tree can attain a height of 16 metres in 
eastern Pondoland and Natal — rather higher than the heights 
recorded by collectors of the species. Wild & Vidigal, l.c., 
incorrectly attribute the combination T. grandifolia to 
Durand & Schinz. 



Aphloia (DC.) Benn. in Benn. & R. Br., PL Jav. Rar. 2: 192 (1840); Benth. & Hook.f., Gen. PI. 1, 
1: 126 (1862); Perrier in FI. Madag. Flacourt. 12 (1946); Wild in F.Z. 1: 279 (1960); Wild & 
Vidigal in FI. Mocamb. Flacourt. 19 (1973). Type species: A. theiformis (Vahl) Benn. 

Prockia sect. Aphloia DC., Prodr. 1: 261 (1824). 

Neumannia A. Rich, in La Sagra, Hist. Fis. Cuba 10: 96 (1843). 

Trees or shrubs. Leaves alternate, simple, petiolate; stipules minute, caducous. Flowers 
bisexual, axillary, solitary, in few-flowered fascicles or racemes. Sepals 4—6, free, except at base, 
orbicular, concave, imbricate, somewhat leathery. Petals 0. Stamens numerous, free, inserted 
outside or on a flat disc. Ovary sessile or shortly stipitate, superior, 1 -locular, with few to several 
ovules on a single parietal placenta; style very short; stigma broadly peltate with a median furrow. 
Fruit a fleshy berry with c. 6 seeds. Seeds obovoid or globose, somewhat flattened, incurved testa 
crustaceous with thin endosperm; embryo incurved; cotyledons ovate. 

About 6 species occurring from Tanzania, southwards to Natal and in Madagascar, the Mascarene and Comoro Islands 
and Seychelles. The generic name is derived from the Greek for “without a corolla (petals) or bark” (Wittstein). 

Aphloia theiformis (Vahl) Benn. in Benn. 
& R. Br., PI. Jav. Rar. 2: 192 (1840); Bak.f. in 
J. Linn. Soc. (Bot.) 40: 23 (191 1); Perrier in FI. 
Madag. Flacourt. 13 (1946); Wild in F.Z. 1: 279 
(1960); Wild & Vidigal in FI. Mocamb. 
Flacourt. 20 (1973). Type: Reunion Island, 
Commerson s.n. (C, holo.; PRE, photo.). 

Lightfootia theiformis Vahl, Symb. Bot. 3: 69 (1794). 
Type as above. 

Prockia theiformis (Vahl) Willd., Sp. PI. 2, 2: 1214 
(1799). Type as above. 

N eumannia theiformis (Vahl) A. Rich, in La Sagra, Hist. 
Fis. Cuba 10: 97 (1845); Dur. & Schinz, Consp. FI. Afr. 1, 
2: 218 (1898); Gilg in Bot. Jahrb. 40: 503 (1908); Pflanzen- 
fam. ed. 2, 21: 437, t.200 (1925); Engl., Pflanzenw. Afr. 3, 



Fig 26—1, Aphloia theiformis, flowering twig, X 1 (Scheepers 679); a, flower, x 5 (Compton s.n.); b. longitudinal 
' section through ovary, x 10 ( Strey 7750); c, fruit, x IVi ( Obermeyer 1107). 



2: 584, t.260 (1921). N. myrtiflora (Galpin) Th. Dur. in 
Dur. & Schinz, Consp. FI. Afr. 1, 2: 218 (1898). Type as for 
Aphloia myrtiflora Galpin. 

Aphloia myrtiflora Galpin in Kew Bull. 1895: 142 
(1895); Burtt Davy, FI. Transv. 1: 215 (1926); Brenan, 
Checklist Tang. Terr. 2: 229 (1949). Type: Transvaal, 
United Ivy Reef, Moodies near Barberton, Galpin 1082 
(PRE, holo. !; BOL!; K!; NH!; SAM!). 

Tree or shrub up to 10 m high. Branchlets 
brown, longitudinally striate, with narrowly 
decurrent wings arising from outer extremities 
of stipular cushions, glabrous. Leaf-blade ellip- 
tic, 3-8 cm long, 1-3 cm wide, apex acuminate, 
acute or rounded, base cuneate to broadly 
cuneate, margin serrulate, often upper 2 / 3 ’s 
only, lateral nerves about 10 pairs, inconspicu- 
ous, glabrous, chartaceous; petiole 2-4 mm 
long. Flowers axillary, solitary or in fascicles 
of 1-3; pedicels greenish, up to 2 cm long, with 
trifid bracteoles up to 1,8 mm long. Sepals 
white, somewhat leathery, broadly elliptic to 
circular, concave, 2, 5-3,5 mm diam., connate 
at base. Stamens: filaments 3 mm long, glab- 
rous; anthers orbicular, 0,7 mm diam. Ovary 
ellipsoid, sometimes shortly stipitate, 3-4 mm 
long; stigma subsessile, peltate, with a median 
groove. Fruit a white, fleshy berry, c. 8 mm 

diam., with persistent stigma on top. Seeds 
several, subglobose-obovoid, somewhat com- 
pressed, 2,5-3 mm diam. Fig. 26. 

A widely distributed species occurring in Tanzania, 
Rhodesia, Mozambique, Transvaal, Swaziland, Natal, 
Madagascar, the Comoro and Mascarene Islands and 

Transvaal. — 2230 (Messina): Entabeni Forest, Galpin 
9558; Hutchinson & Gillett 4226 (K). 2329 (Pietersburg): 
De Hoek, Keet 1185. 2330 (Tzaneen): Westfalia Estate, 
Duiwelskloof, Scheepers 679; Debengeni Falls, Vahr- 
meijer 468. 2430 (Pilgrims Rest): Mariepskop, Van der 
Schijff 6366. 2531 (Komatipoort): Ida Doyer Nature Re- 
serve, Edwards 4124; United Ivy Leaf, Moodies, near 
Barberton, Galpin 1082. 

Swaziland. — 2531 (Komatipoort): Piggs Peak, Comp- 
ton 28228; Kings Bush, Miller 7475. 

Natal. — 2930 (Pietermaritzburg): Zwati, Ndwedwe, 
Strey 7526; 7750a; 7750b. 

A smallish tree or shrub with longitudinally striate 
branchlets, elliptic, serrulate leaves and smallish apetalous 
flowers on longish pedicels. A variable species with numer- 
ous infraspecific taxa described from Madagascar. Perrier 
(l.c. 18) considers the South African plant to be A. 
theiformis (Vahl) Benn. subsp. madagascariensis (Clos.) 
Perrier var. closii Tul. In the present treatment, A. theifor- 
mis is accepted in a broad sense without recognition of 
infraspecific taxa. 



Flacourtia L’Herit., Stirp. Nov. 3: 59, t.30, 30 (3 (1786); Benth. & Hook.f., Gen. PI. 1, 1: 128 
(1862); Sim, For. FI. P.E. Afr. 13 (1909); Engl., Pflanzenw. Afr. 3,2: 585 (1921); Gilg in 
Pflanzenfam. ed. 2,21: 438 (1925); Phill., Gen. ed. 2: 515 (1951); Wild in F.Z. 1: 285 (1960); 
Bamps in F.C.B. Flacourt. 48 (1968); Wild & Vidigal in FI. Mocamb. Flacourt. 27 (1973). Type 
species: F. ramontchi L’Herit. 

Trees or shrubs, often spiny. Leaves alternate, simple, petiolate, mostly crenate. Flowers 
dioecious, rarely bisexual, in short axillary racemes or solitary. Calyx 4— 7-partite or sepals free and 
imbricate. Petals 0. Male flowers: Stamens numerous; filaments terete; anthers small, elliptic or 
somewhat quadrate, sometimes subdidymous. Female flowers: Stamens 0 or few. Ovary 
surrounded by annular or interrupted disc, usually 4— 8-locular with 2 superposed ovules in each 
loculus; styles 4—8, short, thick, more or less radiating, persistent; stigma retuse or marginate. Fruit 
a berry, 5-16-locular. Seed obovoid or subcompressed; testa bony, rough; embryo straight; 
cotyledons flat, ovate; endosperm present. 

About 15 species found in south-east Asia, Malaysia, Polynesia, Madagascar, tropical and southern Africa. The genus 
was named in honour of Etienne de Flacourt (1607-1660), General Director of the French East India Company, Governor of 
Madagascar and author of a history of that island. 

Flacourtica indica (Burm.f.) Merr., In- 
terpr. Rumph. Amboin. 377 (1917); Gilg in 
Pflanzenfam. ed. 2,21: 440, t. 20 1 (1925); Bre- 
nan, Checklist Tang. Terr. 2: 231 (1949); 
Sleumer in FI. Males. Flacourt. 76 (1954); Wild 
in F.Z.: 286 (1960); Bamps in F.C.B. Flacourt. 

48 (1968); Wild & Vidigal in FI. Mocamb. 
Flacourt. 28 (1973). Type: Java, Doery Roekan, 
Burmann collection (G, holo.!; PRE, photo.). 
Gmelina indica Burm.f., FI. Ind. 132, t.39, fig. 5 (1768). 
Flacourtia ramontchi L’Herit., Stirp. Nov. 3: 59, t.30, 30 
(3 (1786); Oliv. in F.T.A. 1: 120 (1868); Sim, For. FI. P.E. 



Fig. 27. — 1, Flacourtia indica, flowering twig, x 1 ( Smuts & Gillett 3415); a, fruit, x 1 (A cocks 12875); b, male flower, X 
6 (Codd & De Winter 5057); c, bisexual flower, X 6 (Codd & De Winter 5057); d, female flower, X 6 (Edwards s.n.); 
e, longitudinal section through $ flower, X 6 (Edwards s.n.). 



Afr. 13 (1909); Bak.f. in J. Linn. Soc. (Bot.) 40: 23 (1911); 
R.E. Fr., Schwed. Rhod.-Kongo-Exped. 1: 157 (1914); 
Perrier, FI. Madag. Flacourt. 9 (1946). Syntypes: Madagas- 
car, Poivre s.n.; Commerson s.n. (P!). F. hirtiuscula Oliv., 
l.c. 121 (1868); Sim, l.c. 13 (1909); Bak. f„ l.c. 23 (1911); 
Gilg, l.c. 440 (1925); Burtt Davy, FI. Transv. 1: 215 (1926). 
Type: Mozambique, near Sena, Kirk s.n. (K, holo. !; PRE, 

Tree or shrub, 1,5-16 m high. Bark 
smooth, stippled white (teste De Winter 7748) 
or rough and yellowish or orange-brown, occa- 
sionally silvery on young branches (Wild l.c.). 
Branches with or without axillary straight 
spines, sometimes with branching spines up to 
12 cm long on the trunk near the base or with 
very spiny coppice shoots, glabrous or pubes- 
cent. Leaf-blade elliptic, ovate or obovate, 
5-9,5 cm long, 3,5-6 cm wide, apex acuminate, 
acute or rounded, base cuneate to rounded, 
margins crenate, crenate-serrate or entire, veins 
4—6 pairs, more prominent on lower surface, 
chartaceous, glabrous to densely pubescent; 
petiole 4-10 mm long, channelled above, 
glabrous or pubescent. Flowers dioecious or 
bisexual, greenish. Calyx segments or sepals 
often unequal, narrowly to broadly ovate, 2-3 
mm long, 1,5-3 mm wide, ciliate. Male flow- 
ers: Stamens very numerous, surrounded by 
disc of fleshy free glands; filaments terete, 2,2 
mm long. Female flowers: Ovary globose c. 3 
mm diam., surrounded by an annular disc; 
styles 4—8, channelled above; stigmas retuse. 

Bisexual flowers as female flowers, but with up 
to 10 stamens. Fruit red, globose, up to 3 cm 
diam., fleshy with persistent styles. Seeds c. 10, 
obovoid, subcompressed, c. 7 mm diam.; testa 
beige, hard, rough. Fig. 27. 

Occurring in scrub, woodland and forest, sometimes 
riverine, in the Transvaal. Also found in south and eastern 
tropical Africa, Madagascar, south-east Asia, Polynesia and 

Transvaal. — 2229 (Waterpoort): northern end of Wyl- 
lies Poort, De Winter 7748. 2230 (Messina): Makonde 
Mission Station, Codd6851. 2328 (Baltimore): 37 km E. of 
Potgietersrust on road to Swartwater, Van der Schijff 5323. 
2329 (Pietersburg): Louis Trichardt, Gerstner 5680. 2330 
(Tzaneen): Elim, Obermeyer 541. 2427 (Thabazimbi): 
upper slopes of Krantzberg, Prosser 1717 (K). 2428 
(Nylstroom): between Nylstroom and Warmbaths, Pole 
Evans 188. 2429 (Zebediela): Potgietersrust to Zebediela, 
Pole Evans 3104 (23). 2431 (Acomhoek): 5 km N.E. of 
Skukuza on Tshokwane Road, Codd & De Winter 5057. 
2526 (Zeerust): Enselberg camp, Pole Evans 2225. 2527 
(Rustenburg): Jacksontuin, Brits, Mogg 14979. 2529 (Wit- 
bank): Loskop Dam, Theron 923. 2530 (Lydenburg): Low- 
veld Botanic Garden, Buitendag 815. 2531 (Komatipoort): 
Pretoriuskop, Van der Schijff 3924. 

A small tree with or without spines, elliptic, ovate or 
obovate leaves with usually crenate margins and red, 
globose fruits up to 3 cm in diam. An extremely variable 
species as regards leaf shape and degree of pubescence. The 
plant is variously known as the Madagascar, Mauritius, 
Governor’s or Batoko Plum and is cultivated in the tropics 
of both the Old and New Worlds, because of its edible fruit. 
The rather acid fruits may be eaten raw or cooked to make 
jam and preserves. The wood is used for making agricul- 
tural implements and the plant is used medicinally in 
tropical Africa. 

5328 12. DOVYALIS* 

Dovyalis E. Mey. ex Am. in Hook., J. Bot. 3: 251 (1841); Drege, Cat. PI. Afr. Austr. 3 (1840); 
Zwei Pfi. Doc. 125 (1843), p. 180, as Doryalis, nom. nud.; Endl., Gen. PI. Suppl. 2: 91 (1842); 
Sond. in Linnaea 23; 12 (1850); Harv. in F.C. 1: 69 (1860); Benth. & Hook.f., Gen. PI. 1: 128 
(1862); Harv., Gen. PI. ed. 2; 16 (1868); Warb. in Pflanzenfam. 3,6a: 44 (1893), as Doryalis-, Sim, 
For. FI. Cape Col. 129 (1907); For. FI. P.E. Afr. 13 (1909); Engl, in Bot. Jahrb. 40: 504 (1908), as 
Doryalis', Gilg in Pflanzenfam. ed. 2, 21: 440 (1925); Burtt Davy, FI. Transv. 1: 216 (1926); Phill., 
Gen. ed. 2: 515 (1951); Wild in F.Z. 1: 281 (1960); Sleumer in Bot. Jahrb. 92: 64 (1972). Type 
species: D. zizyphoides E. Mey. ex Arn. 

Aberia Hochst. in Flora 27: 2 (1844); Sond. in Linnaea 23: 9 (1850); Harv. in F.C. 1: 70 (1860); in F.C. 2: Addenda 584 
(1862); Benth. & Hook.f., Gen. PI. 1: 128 (1862); Oliv. in F.T.A. 1: 121 (1868); Harv., Gen. PI. ed. 2: 16 (1868). Type 
species: A. verrucosa Hochst. 

Trees or shrubs, evergreen or deciduous, dioecious or rarely polygamous, often armed with 
spines; branches lenticellate. Leaves exstipulate, alternate or sometimes fascicled, petiolate; blade 
3-5-veined at or near the base, margins entire, crenate or serrate. Male flowers pedicellate, in 
fascicles of 2-10, or 5-10 flowers in groups of 3; pedicels surrounded at the base by small hairy 

By J. E. Langenegger. 



scales; calyx (tepals of Sleumer l.c.) 2-6-lobed; corolla 0; stamens 10-many; filaments surrounded 
at the base by sparsely to densely hairy nectaries which form a honeycomb structure; anthers 
bilocular, dehiscing by means of longitudinal slits. Female flowers pedicellate, axillary, solitary or 
in fascicles of 2 or 3; pedicels surrounded at the base by small hairy scale-like bracts; calyx (tepals) 
deeply 3-7-lobed; lobes sometimes revolute and sometimes fringed with stipitate glands; corolla 0; 
disc annular, lobed, glutinous, hairy, sparsely hairy or glabrous; ovary unilocular or sometimes 
incompletely 2 or 3-locular; placentas 2-7, parietal, each placenta with 1 or 2 ovules; styles 
channelled, as many as placentas. Fruit a subglobose to oblong berry, minutely papillose, glabrous 
or hairy. Seeds embedded in fleshy pulp, elliptic to broadly elliptic, glabrous to densely woolly; 
testa leathery; embryo straight; cotyledons flat, elliptic, sometimes thin and pinnately veined; 
endosperm present. 

A genus of about 20 species, widespread in Africa and extending to Sri Lanka and New Guinea; 6 species recognized in 
Southern Africa. 

The spelling of the generic name has been the subject of some confusion. It first appeared as a nomen nudum in Drege, 
Cat. PI. Afr. Austr. (1840), spelt Dovyalis E. Mey. A year later, in 1841, Arnott provided a description of Dovyalis. In 
Drege's Zwei Pfi. Doc. (1843) the generic name was spelt Dovyalis in the text and Doryalis in the index. It seems very 
probable that the latter spelling was intended, because (a) it is placed in the correct alphabetical sequence in the index, and 
(b) it is no doubt derived from the Greek “dory”, a spear, in allusion to the conspicuous spines found on most plants. 
However, although Warburg in 1895, restored Doryalis, the original spelling Dovyalis used by Amott must be adopted 
according to Article 73 of the International Code of Botanical Nomenclature (1972). 

Ovary with 5-7 placentas, each placenta with 2 ovules; styles 5-7; fruit with many seeds; fruit 3-4 cm in 
diam.; ultimate branchlets often abbreviated, up to about 2 cm long or forming cushion-like structures; leaves 

fascicled or, on young shoots, alternate 1. D. caffra 

Ovary with 2 or 3 placentas, each placenta with 1 ovule; styles 2 or 3; fruit with 1-3 seeds; fruit 0,6-2 cm in 
diam.; ultimate branchlets often abbreviated, but not forming cushion-like structures; leaves not 

Exocarp of fruit glabrous: 

Testa of seed glabrous 2. D. rotundifolia 

Testa of seed hairy: 

Leaves ovate to elliptic-ovate; leaf base cordate to truncate; the secondary nerves tending to 

spread towards the leaf margin 3. D. rhamnoides 

Leaves obovate; leaf base cuneate; the secondary nerves tending to ascend to the leaf 

tip 4. D. longispina 

Exocarp of fruit hairy or papillose: 

Calyx (tepal) lobes of ? flowers 4-6 mm long in anthesis, accrescent 5. D. zeyheri 

Calyx (tepal) lobes of ? flowers 1,5-2 mm long in anthesis, not accrescent 6. D. lucida 

1. Dovyalis caffra (Hook.f. & Harv.) 
Hook.f in Harv., Gen. PI. ed. 2; 16 (1868); 
Warb. in Pflanzenfam. 3, 6a: 44 (1893), as 
Doryalis ; Sim, For. FI. Cape Col. 129 (1907); 
For.' FI. P. E. Afr. 13 (1909); Engl, in Bot. 
Jahrb. 40: 604 (1908), as Doryalis ; Marloth, FI. 
S. Afr. 2: 195 (1925); Gilg in Pflanzenfam. ed. 
2, 21: 440 (1925); Burtt Davy, FI. Transv. 1: 
216 (1926); Wild in F.Z. 1: 285 (1960); 
Sleumer in Bot. Jahrb. 92: 68 (1972). Syntypes: 
Cape, eastern districts, R. Hallack s.n. (TCD!; 
PRE, photo.); Kaffirland, Holland s.n. 

Aberia caffra Hook.f. & Harv. in F.C. 2: 584 (1862); T. 
Anders, in J. Linn. Soc. 7: 68 (1862); Fourcade, Rep. Natal 
Forests 94 (1889). A. edulis T. Anders, l.c. 68, nom. nud. 

Tree or shrub up to 6 m tall, evergreen, 
dioecious or rarely polygamous, armed with 
spines up to 6 cm long; bark ashen-white to 
brown with longitudinal ridges and fissures; 
ultimate branchlets often much abbreviated, up 
to about 2 cm long or forming cushion-like 
structures. Leaves fascicled or, on young 
shoots, alternate; blade soft to coriaceous, obo- 
vate or sometimes elliptic-rhomboid, 2-5,5 cm 
long, 1-3 cm broad, glabrous or rarely puberul- 
ous, with prominent venation on both sides, 
3-5-veined from the base, apex acute to obtuse 
or emarginate, base usually cuneate, margin 
entire, slightly revolute; petiole 2-4 mm long, 
glabrous. Male flowers light green, fascicled. 



with 5-10 flowers in groups of 3; calyx 2-5- 
lobed; lobes 3 mm long, elliptic, pubescent; 
stamens numerous; filaments 5 mm long; nec- 
taries hairy. F emale flowers light green, solitary 
or in fascicles of 2-3 on abbreviated shoots; 
pedicels 4—10 mm long; calyx deeply 5-7- 
lobed, lobes 3 mm long, elliptic-lanceolate, 
puberulous, acuminate, somewhat recurved; 
disc sparsely hairy; ovary 5-7-lobed, unilocu- 
lar; placentas 5-7, each placenta with 2 ovules; 
styles 5-7. Fruit subglobose, 3-6 cm in diam., 
minutely velvety, bright yellow. Seeds many, 
woolly, 1 cm long. Fig. 28: 2; 29: 1. 

A common constituent of mixed scrub, riverine bush, 
open woodland and forest; distributed on a variety of soils 
from the Bathurst district in the eastern Cape northwards 
along the coast and in river valleys to Natal, eastern, central 
and northern Transvaal, and extending to Rhodesia. It has 
been introduced into California and Australia. 

Transvaal. — 2231 (Pafuri): Shipudza area, Punda 
Milia, Van Wyk 4741. 2329 (Pietersburg): on tropic of 
Capricorn, Story 4840. 2330 (Tzaneen): Elim, Obermeyer 
752 . 2430 (Pilgrims Rest): Ohrigstad, Burn Davy 7314; 
Nooitgedacht Mountain, Young A 657. 2531 

(Komatipoort): Pretoriuskop, Van der Schijff 3902. 

Natal. — 2732 (Ubombo): Mkuze Game Reserve, Ward 
3831. 2830 (Dundee): Keats Drift road from Muden, 
Edwards 913. 2831 (Nkandla): Eshowe, Lawn 1270 (NH); 
Hlabisa, Ward 4783. 

Cape. — 3129 (Port St. Johns): Tabankulu, 6 km from 
Welsh Bridge on Ceveraland road, Story 4218. 3228 
(Butterworth): Kei River Valley, Flanagan 467. 

Dovyalis caffra, the Kei-apple, is an attractive, usually 
evergreen tree which, from December to about February, is 
covered with masses of yellow apricot-like fruits. The fruits 
are edible and are used to make jelly and other preserves. To 
counter the rather acidic taste of the fruits due to a high 
malic acid content, the fruits are often mixed with a sweet 
fruit, e.g. grapes. The plant is spinous and makes an 
excellent impenetrable hedge. It has the added virtue that it 
is fairly drought resistant. 

2. Dovyalis rotundifolia (Thunb.) 
Thunb. & Harv. in F.C. 1: 70 (1860); Warb. in 
Pflanzenfam. 3, 6a: 44 (1893), as Doryalis\ 
Sim, For. FI. Cape Col. 132 (1907); Gilg in Bot. 
Jahrb. 40: 504 (1908); Pflanzenfam. 2, 21: 440 
(1925); Burtt Davy, FI. Transv. 1: 216 (1926); 
Sleumer in Bot. Jahrb. 92: 82 (1972). Type: 
Cape, Thunberg s.n. (UPS, holo.). 

Celastrus rotundifolius Thunb., Prodr. 42 (1794); Willd., 
Sp. PI. 1: 1129 (1798); Thunb., FI. Cap. ed. Schult. 221 

Prockia rotundifolia (Thunb.) Eckl. & Zeyh., Enum. 16 

Dovyalis celastroides Sond. in Linnaea 23: 9 (1850), 
nom. illegit.; Sim, For. FI. P.E. Afr. 13 (1909); Marloth, FI. 
S. Afr. 2: 194 ( 1925); Henkel, Woody PI. Natal 102(1934). 

Tree or shrub up to 6 m tall, dioecious, 
armed with spines up to 8 cm long; bark 
ashen-white with longitudinal ridges and 
fissures, strongly lenticellate. Leaves alternate; 
blade coriaceous, sub-orbicular to obovate, 
2,2-3 cm long, 1,5-2 cm broad, glabrous, with 
prominent venation on both sides, 3-5-veined 
from the base, apex obtuse to emarginate, base 
obtuse to cuneate, margin entire; petiole 2-3 
mm long, glabrous. Male flowers light green, in 
fascicles of 2-10; calyx 4— 6-lobed; lobes 1,5— 
2,5 mm long, lanceolate to obovate-rhomboid, 
pubescent; stamens 9-20, filaments 3 mm long; 
nectaries hairy. Female flowers yellow-green, 
solitary or in fascicles of 2 or 3; pedicels 
0,5-1, 5 mm long; calyx 4— 6-lobed; lobes 1-2 
mm long, obovate, pubescent, acuminate, 
somewhat recurved; disc sparsely hairy; ovary 
2-lobed, unilocular; placentas 2, each placenta 
with 1 ovule; style 2. Fruit oblong, 0,6-1 cm in 
diam., glabrous, red when ripe. Seeds 2, glabr- 
ous, 0,2-0, 6 mm long. Fig. 29: 4. 

Found in coastal bush on sand dunes in the eastern 
Cape Province from Port Elizabeth to East London. 

Cape. — 3227 (Stutterheim): Bonza Bay, Comins 1068. 
3325 (Port Elizabeth): Amanzi, Wells 3178. 3326 

(Grahamstown): Port Alfred, Tyson sub TRV 17246. 3327 
(Peddie): Great Fish River, MacOwan 825 (NBG). 

3. Dovyalis rhamnoides (Burch, ex DC.) 
Burch. & Harv. in F.C. 1: 69 (1860); Fourcade, 
Rep. Natal Forests 94 (1889); Warb. in Pflan- 
zenfam. 3, 6a: 44 (1895), as Doryalis\ Sim, 
For. FI. Cape Col. 131 (1907); Engl, in Bot. 
Jahrb. 40: 504 (1908), as Doryalis; Gilg in 
Pflanzenfam. ed. 2, 21: 440 (1925); Burtt Davy, 
FI. Transv. 1: 216 (1926); Verdoorn, Edible 
Wild Fr. Transv. 33 (1938); Sleumer in Bot. 
Jahrb. 92: 80 (1972). Type: Cape, George, 
Uitenhage and Albany, Burchell 6012 (PRE!). 

Flacourtia rhamnoides Burch, ex DC., Prodr. 1: 256 
(1824); Eckl. & Zeyh., Enum. 16 (1834). 

Dovyalis zizyphoides E. Mey. ex Sond. in Linnaea 23: 12 
(1850); E. Mey. in Drege, Cat. PI. Afr. Austr. 3 (1840), 
nom. nud.; E. Mey. ex Am. in Hook., J. Bot. 3: 251 (1841), 
comb, superfl.; Pappe, FI. Cap. Med. Prodr. 4 (1859); 
Marloth, FI. S. Afr. 2: 194 (1925). Syntypes: Cape, Ruig- 
tevalei, Drege s.n. (PRE!; S!); Krakakamma and 
Olifantshoek, Ecklon 115 (S ! ; PRE, photo.); Van Stadens 
Mts, Zeyher s.n. (S!; PRE, photo.); Port Natal, Gueinzius 
58 (S!; PRE, photo.). 

Tree or shrub up to 5 m tall, dioecious, 
armed with spines up to 6,2 cm long; bark white 
to brown with shallow longitudinal ridges and 
fissures, young twigs and spines pubescent. 



Fig. 28. — 1, Dovyalis rhamnoides, branch with male flowers, x % (Wells 3912). 2, D. caffra, branch with female 
flowers, X 2 h; 2a, female flowers, x 573; 2b, cross-section of ovary, x 573; 2c, male flower, X 6; 2d, stamen, X 5 V 3 
( Thom 1 and 2). 3, D. longispina, longitudinal section of female flower, x 573; 3a, cross-section of ovary, x 573 
( Bourquin 557). 



Leaves alternate; blade thin and somewhat 
membranous, ovate to elliptic-ovate, 2-4,5 cm 
long, 1-2,5 cm broad, glabrous or sometimes 
puberulous especially on the veins, veins prom- 
inent on both surfaces, 3-5-veined from the 
base, apex acute to obtuse, base truncate to 
cordate or sometimes cuneate, margin entire to 
crenate; petiole 2-3 mm long, glabrous or 
puberulous. Male flowers light green, in fasci- 
cles of 3-6; calyx 4— 5-lobed, lobes 2-3 mm 
long, ovate, pubescent; stamens 15, filaments 
1,5 mm long; nectaries hairy. Female flowers 
light green, solitary or in fascicles of 2 or 3; 
pedicels 2-3 mm long; calyx deeply 5-lobed, 
lobes 1-5 mm long, linear, acuminate, erect, 
puberulous and usually fringed with stipitate 
glands; disc hairy; ovary 2-lobed, unilocular, 
with 2 much-reduced placentas, each placenta 
with 1 ovule; styles 2. Fruit ellipsoid to ovoid, 

1 cm in diam., glabrous, orange to scarlet red 
when ripe. Seeds 2-3, 9 mm long, slightly 
woolly. Fig. 28: 1; 29: 5. 

Found in dune forests, on riverbanks, in temperate 
ravine forest, scrub forest and thomveld. Recorded from the 
eastern Cape, Natal, Zululand, Swaziland and the Trans- 

Transvaal. — 2431 (Acomhoek): Lothian, Strey 3555. 

Swaziland. — 2631 (Mbabane): Hlatikulu, Compton 

Natal. — 2731 (Louwsburg): Wendelane Kloof, 
Gerstner 4674. 2831 (Nkandla): Mtunzini, Edwards 1582. 
2832 (Mtubatuba): Hluhluwe Game Reserve, Ward 1690. 
2930 (Pietermaritzburg): Mt. Ashley, Lions River, Moll 
1308. 2931 (Stanger): Groutville, A cocks 11820. 3030 (Port 
Shepstone): Umpanyoni River, Rudatis 1989 (STE). 

Cape. — 3227 (Stutterheim): Stutterheim Commonage, 
Acocks 9720. 3228 (Butterworth): Komga, Flanagan 374 
(NBG). 3322 (Beaufort West): Homtini River gorge, Taylor 

s. n. (Saasveld Herb.). 3325 (Port Elizabeth): Van Staadens 
Kloof, Taylor 834. 3326 (Fort Beaufort): Kowie, Britten 
768. 3422 (Mossel Bay): Gouwkamma River, Keet 611 
(STE). 3423 (Knysna): Knysna River, Schonland 3431 
(GRA). 3424 (Humansdorp): Humansdorp, Galpin 4577. 

According to Smith (1966), D. rhamnoides was known 
as Wynbessie, because of the pleasantly vinous taste of the 
ripe fruits from which brandy and vinegar were made in the 
early settler days of the eastern Cape. The wood of this 
species was formerly used for making yokes, wagons and 
agricultural implements. 

4. Dovyalis longispina (Harv.) Warb. in 
Pflanzenfam. 3, 6a: 44 (1893), as Doryalis-, 
Gilg in Bot. Jahrb. 40: 506 (1908), as Doryalis-, 
Pflanzenfam. ed. 2, 21: 441 (1925). Syntypes: 
Natal, near Durban, Gerrard & McKen 542; 
541 (TCD, holo. !; PRE, photo.). 

Aberia? longispina Harv. in F.C. 2, App. : 585 (1862). 

Dovyalis celastroides sensu Sim, For. FI. P.E. Afr. 13, 

t. 17fB (1909). 

Tree or shrub up to 1 1,5 m high, dioecious, 
armed with spines up to 8 cm long; bark 
ashen-white with longitudinal ridges and 
fissures, strongly lenticellate. Leaves alternate; 
blade soft to coriaceous, obovate to rhomboid- 
elliptic, seldom orbicular, 2, 2-9, 5 cm long, 
1,5-5, 5 cm broad, glabrous, sometimes 
puberulous on veins, venation very prominent 
on both sides, 3-5-veined from the base, apex 
acute to emarginate, base cuneate to obtuse, 
margin entire to sub-undulate; petiole 2-6 mm 
long, glabrous. Male flowers light green, in 
fascicles of 2-10; calyx 4— 6-lobed; lobes 1,5— 
2,5 mm long, obovate-rhomboid, pubescent; 
stamens 9-20; filaments 3 mm long; nectaries 
hairy. F emale flowers yellow-green, solitary or 
in fascicles of 2 or 3; pedicel 3-5 mm long; 
calyx 4— 6-lobed; lobes 3-5 mm long, elliptic- 
lanceolate to obovate, pubescent, sometimes 
fringed with gland-tipped segments, acute, erect 
or somewhat recurved; disc hairy; ovary 
2-lobed, unilocular, placentas 2, each with 1 
ovule; styles 2. Fruit oblong, 1,5 cm in diam., 
glabrous, orange or red with white spots when 
ripe. Seeds 2, woolly, 0,9 cm long. Fig. 28: 3; 
29: 2. 

Found in coastal bush on sand dunes and in low-lying 
areas near mangroves. Further inland, it occurs in mixed 
scrub and scrub forest, on sandy soil. Recorded from the 
northern coastal areas of Natal, northwards to Mozambique. 

Natal. — 2732 (Ubombo): between Sordwana Bay and 
Jozini, Van der Schijff 6577. 2831 (Nkandla): Umlalazi 
Nature Reserve, Fakude 18. 2832 (Mtubatuba): Hluhluwe 
Game Reserve, Ward 4782. 2930 (Pietermaritzburg): near 
Isipingo River, Ward 493. 2931 (Stanger): Hlogweni 
Forest, Moll 3621. 

5. Dovyalis zeyheri (Sond.) Warb. in 
Pflanzenfam. 3, 6a: 44 (1893), as Doryalis-, 
Gilg in Bot. Jahrb. 40: 506 (1908), as Doryalis-, 
Pflanzenfam. ed. 2, 21: 440 (1925); Burtt Davy, 
FI. Transv. 1: 216 (1926); Wild in F.Z. 1: 284 
(1960); Killick in Flow. PI. Afr. 39: 1. 1546 
(1969); Sleumer in Bot. Jahrb. 92: 84 (1972). 
Type: Transvaal, Crocodile River, Zeyher 52 
(S, holo.!; TCD!; PRE, photo.). 

Aberia zeyheri Sond. in Linnaea 23: 10 (1850); Harv. in 
F.C. 1: 70 (1860); Szyszyl., Polypet. Thalam. Rehm. 112 
(1888). — var. velutina Szyszyl., l.c. (1888). Syntypes: 
Transvaal, Wonderboompoort, Rehmann 4557 (Bf, Z); 
Aapiespoort, Rehmann 4097 (Bf; BM; K, lecto.); Rehmann 
4098 (Bf; Z). A. tristis Sond. in Linnaea 23: 9 (1850); 
Harv. in F.C. 1: 70 (1860). Type: Cape, Phillipstown, Kat 
River, Ecklon <6 Zeyher s.n. (S, holo.!; SAM!; PRE, 

Dovyalis tristis (Sond.) Warb. in Pflanzenfam. 3, 6a: 44 
(1893), as Doryalis ; Sim, For. FI. Cape Col. 129 (1907); 
Gilg in Bot. Jahrb. 40: 506 (1908), as Doryalis ; Pflanzen- 



Fig. 29. — 1, Dovyalis caffra, leaf, X 2 A; la, fruit, X 2 h; lb, seed, x l l /3 (Edwards 5328). 2, D. longispina, fruiting twig, 
x 2 / 3 ; 2a, seed, X IV3 (De Winter & Vahrmeijer 8426). 3, D. zeyheri, fruiting twig, x 2 /y, 3a, seed, x IV3 (Letty 
477). 4, D. rotundifolia, leaf, X 2 h \ 4a, fruit, X 2 h\ 4b, seed, x IV3 (Tyson 186).. 5, D. rhamnoides, leaf, X 2 ly, 5a, 
fruit, X 5b, seed, X IV3 (De Winter 7650). 6, D. lucida, leaf, X 2 h; 6a, fruit, X 2 /y, 6b, seed, x IV3 (Forester, 
Entabeni, December 1969). 



fam. ed. 2, 21: 441 (1925); Marloth, FI. S. Afr. 2: 194 
(1925); Burtt Davy, FI. Transv. 1: 216 (1926). — var. 
depauperata Sim, For. FI. Cape Col. 131 (1907). Type: 
Cape, Keiskama, about Line Drift, Sim 2570 (NU, holo. !). 
D. revoluta Thom in Bothalia 10: 434 (1971). Type: Natal, 
False Bay, Moll 5112 (PRE!). 

Tree or shrub up to 11 m tall, dioecious, 
often armed with pubescent or glabrous spines 
up to 3,5 cm long; bark ashen-white with 
longitudinal ridges and shallow fissures; ulti- 
mate branchlets often abbreviated; young twigs 
pubescent. Leaves alternate; blade thin and soft, 
sometimes coriaceous when mature, obovate to 
rhomboid-elliptic, 1,5-6 cm long, 1-4 cm 
broad, pubescent to glabrescent, 3-veined from 
the base, apex acute to obtuse, or emarginate, 
base cuneate, margin entire to crenate; petiole 

2- 5 mm long, pubescent or sometimes glabr- 
ous. Male flowers yellow-green, in fascicles of 

3- 6; calyx 4— 6-lobed, lobes 4 mm long, nar- 
rowly ovate, pubescent; stamens 24—48, fila- 
ments 2,5-3 mm long; nectaries densely hairy. 
Female flowers yellow-green, solitary or rarely 
in pairs; pedicels 4—7 mm long; calyx deeply 

4- 7-lobed; lobes 4—6 mm long, lanceolate- 
obovate, acute, tomentose, somewhat recurved; 
disc densely hairy; ovary 2 or 3-lobed, almost 
completely 2 or 3-locular, placentas 2 or 3, each 
placenta with 1 ovule; style 2-3. Fruit oblong, 
1, 1-1,5 cm in diam., densely and shortly hairy, 
orange to red when ripe. Seeds 2-3, hairy, 7 
mm long, with a coffee smell. Fig. 29; 3. 

Occurs in several veld types including forest margins, 
open woodland and streambanks, often among rocks on a 
variety of soil types. Recorded from the coastal regions of 
the eastern Cape, inland to as far as Queenstown, extending 
to Natal, Zululand, Swaziland, and the southern, eastern and 
northern Transvaal. Also recorded from southern Rhodesia. 

Transvaal. — 2230 (Messina): Makonde Mission Sta- 
tion, Codd6817. 2328 (Baltimore): Leipzig, Brcmekamp & 
Schweickerdt 56. 2329 (Pietersburg): Dorps River, Louis 
Trichardt, Gerstner 5697. 2330 (Tzaneen): Letaba, Scheep- 
ers 999. 2429 (Zebediela): Farm Magnet Fleights, Barnard 
& Mogg 864. 2430 (Pilgrims Rest): Sekukuniland, Barnard 
423. 2527 (Rustenburg): Hartebeespoortdam, Vahrmeijer 
413. 2528 (Pretoria): Liebenberg 3. 2529 (Witbank): Los- 
kopdam Nature Reserve, Mogg 30385. 2531 

(Komatipoort): Barberton, Pott 17749. 2627 (Potch- 
efstroom): Witpoortjie, Rogers 18538. 

Swaziland. — 2631 (Mbabane): Tulwane, Compton 

Natal.— 2731 (Louwsburg): 21 km N. of Nongoma, 
Acocks 13020. 2829 (Harrismith): Collings Pass, Acocks 
13859. 2830 (Dundee): Roodedraai, Acocks 10482. 2832 
(Mtubatuba): False Bay, Edwards 3199; Moll 5109; 5112. 
2929 (Underberg): Cathedral Peak, Killick 1901. 2930 
(Pietermaritzburg): Albert Falls, Comins 466. 

Cape. — 3128 (Umtata): Mqanduli, Pegler 592. 3225 
(Somerset East): Bosberg, Van der Walt 354. 3226 (Fort 

Beaufort): Kei River Valley, Galpin 8074. 3227 (Stut- 
terheim): Cata Forest, Wells 3229. 3327 (Peddie): Sim 

The species varies a great deal in its pubescence. The 
unarmed form with relatively glabrous leaves has in the past 
been separated as D. tristis, while D. zeyheri is described as 
being spiny and having pubescent leaves. Intermediates are, 
however, so common that the step taken by Wild (l.c.) to 
merge the two species is supported. Following Sleumer 
(l.c.), D. revoluta is treated here as a local form of D. 
zeyheri with glabrous leaves and papillose-puberulous 
ovaries and fruit. 

6. Dovyalis lucida Sim, For. FI. Cape 
Col. 131, t.6 (1907); Wild in F.Z. 1: 282 
(1960); Sleumer in Bot. Jahrb. 92: 87 (1972). 
Type: Eastern Cape, Plate 6, For. FI. Cape Col. 
(1907), no specimen found. 

Tree or shrub up to 10 m tall, dioecious, 
rarely armed with spines up to 3,5 cm long, 
bark yellow- white to brown, strongly lenticel- 
late. Leaves alternate; blade soft to coriaceous, 
obovate to rhomboid, rarely ovate, 2-7,5 cm 
long, 1,2-4 cm broad, glabrous, shining and 
dark green above, paler and dull below, veins 
prominent on both sides, 3-5-veined from the 
base; apex acuminate, rarely emarginate, base 
cuneate, rarely obtuse, margin entire or serrate; 
petiole 3-5 mm long, glabrous. Male flowers 
light green, in fascicles of 3-10, rarely in a 
small raceme; pedicels 1-3 mm long; calyx 
3-5-lobed, lobes 3 mm long, ovate-elliptic, 
pubescent; stamens 1 1-27, filaments 3 mm 
long; nectaries densely hairy. Female flowers 
yellow-green, solitary or in fascicles of 2 or 3; 
pedicels 3-4 mm long; calyx 3-5-lobed, lobes 
1,5-2 mm long, obovate, obtuse, pubescent, 
somewhat recurved; disc densely hairy; ovary 2 
or 3-lobed, unilocular, with 2 or 3 placentas, 
each placenta with 1 ovule; styles 2 or 3. Fruit 
ellipsoid, 1,5 cm in diam., minutely papillose, 
orange to red when ripe. Seeds 2 or 3, hairy, 1,1 
cm long. Fig. 29: 6. 

Found in scrub forest and forest from Port Elizabeth in 
the Eastern Cape, northwards through Natal to the Sout- 
pansberg in the Northern Transvaal, extending to the eastern 
districts of Rhodesia. 

Transvaal. — 2230 (Messina): Entabeni Forest Reserve, 
Leistner, Thom & Gillham 3310. 2329 (Pietersburg): Farm 
Seodin, Leistner, Thom & Gillham 3279. 2330 (Tzaneen): 
Magoebaskloof, Codd 1687. 2430 (Pilgrims Rest): 1,6 km 
S. of Graskop, Leistner, Thom & Gillham 3322. 

Natal. — 2830 (Dundee): Qudeni Forest, Edwards 2633. 
2831 (Nkandla): Inhlazatje Mt., Acocks 11686. 2929 (Un- 
derberg): Ntambamhlope, Edwards 695. 2930 (Pieter- 
maritzburg): Umgeni Poort Forest, Moll 861. 

Cape. — 3128 (Umtata): Engcobo, Acocks 13815. 3226 
(Fort Beaufort): Hogsback, Story 369. 3227 (Stutterheim): 
King William’s Town, Comins 1024. 3325 (Port Elizabeth): 
Van Stadens Kloof, Taylor 837. 3326 (Grahamstown): 
Albany, Wells 3016. 





Casearia Jacq., Enum. Syst. PI. Ins. Carib. 4, 21 (1760); Benth. & Hook.f., Gen. PI. 1,3: 796 
(1867); Gilg in Bot. Jahrb. 40: 510 (1908); Engl., Pflanzenw. Afr. 3,2: 589 (1921); Gilg in 
Pflanzenfam. ed. 2, 21: 451 (1925); Phill., Gen. ed. 2: 516 (1951); Wild in F.Z. 1: 293 (1960); 
Sleumer in Bull. Jard. Bot. Nat. Belg. 41: 397 (1971); Wild & Vidigal in FI. Mocamb. Flacourt. 35 
(1973). Type species: C. nitida (L.) Jacq. 

Trees or shrubs. Leaves alternate, simple, petiolate, entire, serrate or with subspinose teeth; 
stipules small. Inflorescence an umbel or flowers fascicled or rarely solitary. Flowers bisexual. 
Calyx-tube short or subelongated or 0; sepals 4—6, suborbicular, concave, imbricate. Petals 0. 
Stamens 6-15, rarely 20-40, free or connate, hypogynous or inserted on the calyx-tube or at the 
base of calyx; filaments linear, connective of anthers sometimes penicillate at the apex; staminodes 
alternating with the stamens, sometimes oblong and hairy at the apex. Ovary superior, 1 -locular 
with few to many ovules on 3 or 4 parietal placentas; style short; stigma capitate or peltate or 
stigmas 3. Fruit a subfleshy or dry capsule, opening by 3 or 4 valves, few to many-seeded. Seeds 
sometimes angled, with a fleshy aril; testa chartaceous; embryo straight; cotyledons flat, elliptic or 
suborbicular; endosperm present. 

Species about 160 in tropical countries of the world. Casearia was named in honour of Johann Casearius, a Dutch 
missionary in Indo-China in the 1700’s, who was responsible for the first part of the Hortus Malabaricus (Wittstein). 

Casearia gladiiformis Mast, in F.T.A. 2: 
493 (1871); Gilg in Bot. Jahrb. 40: 510 (1908); 
Engl., Pflanzenw. Afr. 3, 2: 589 (1921); Gilg in 
Pflanzenfam. ed. 2, 21: 454 (1925); Wild in 
F.Z. 1: 293 (1960); Sleumer in Bull. Jard. Bot. 
Nat. Belg. 41: 423 (1971); Wild & Vidigal in 
FI. Mocamb. Flacourt. 35 (1973). Type: 
Mozambique, Shupanga, Zambesi River, Kirk 
s.n. (K, holo. !; PRE, photo.). 

C. junodii Schinz in Mem. Herb. Boiss. 10: 52 (1900); 
Gilg in Bot. Jahrb. 40: 513 (1908); Engl., Pflanzenw. Afr. 3, 
2: 590 (1921); Gilg in Pflanzenfam. ed. 2, 21: 454 (1925); 
Henkel, Woody PI. Natal 87 (1934). Type: Mozambique, 
Delagoa Bay, Junod 351 (Z, holo.!; K!; PRE, photo.). 

Tree up to 25 m high; young branches 
puberulous, becoming glabrous; bark grey, 
smooth. Leaf -blade elliptic or ovate, 7-18 cm 
long, 3-7 cm wide, apex acuminate, acute or 
obtuse, cuneate and oblique at base, margins 
usually entire but sometimes serrate-crenate 
especially young and coppice shoot leaves, 
coriaceous, glabrous or puberulous on midrib 
when young, with circular and linear pellucid 
dots, lateral nerves 7-9, V 2 -angled, curved, 
more prominent on lower than upper side; 
petiole 0, 1-2 cm long, channelled above, 
puberulous or glabrous, dark green and shiny. 
Flowers greenish, in dense, axillary fascicles on 
a cushion of minute, brown, membranous, 
ovate bracteoles with ciliate margins; pedicels 
1-3 mm long, puberulous. Sepals 5, broadly 
elliptic, 3 mm long, 2 mm wide, puberulous 
dorsally. Staminal tube united for about 0,3-0, 5 
mm and then dividing into 10 fertile stamens 

and 10 alternating staminodes; filaments 1 mm 
long, glabrous; anthers ovate, 0,8 mm long, 
apiculate; staminodes 1 mm long, villous at 
apex. Ovary ovoid, 1,75 mm long, 1 mm wide, 
glabrous or with long hairs in upper V 3 ; style 
0,5 mm long; stigma capitate. Fruit yellowish- 
orange, angular, ellipsoid capsule, splitting 
from apex into 3-4 longitudinal valves. Seeds 
c. 10, ellipsoid or ovoid, 4,5 mm long, 2 mm 
wide, with pale beige testa clasped by fleshy 
aril. Fig. 30. 

Occurring in forest, chiefly along the coast, in Natal 
and the eastern Cape. Also found in Kenya, Tanzania, 
Malawi and Mozambique. 

Natal. — 2632 (Bela Vista): Mangusi Forest, Venter 
6220; Lake Nhlange, Kosi System, Vahrmeijer & Tolken 
953. 2732 (Ubombo): Lake Sibayi, Stephen 461. 2831 
(Nkandla): Lower Nkandla Forest, Edwards 1446. 2832 
(Mtubatuba): Charter’s Creek, Ward 3029. 2930 (Pieter- 
maritzburg): Howick, Cooper 3; Farm “The Start”, Lions 
River, Moll 3366. 3029 (Kokstad): Mt. Ingeli, Nicholson 
760. 3030 (Port Shepstone): Paddock, Strey 5995. 

Cape. — 3128 (Umtata): Mpane Forest, Elliotdale, Miller 
sub PRF 5824. 3129 (Port St. Johns): Qokama, Ngqeleni 
District, Acocks 12820; Egossa Forest, Strey 8870. 3228 
(Butterworth): Manubi Forest, Kentani, Marais 481. 

A tree with elliptic or ovate leaves varying considera- 
bly in size with margins usually entire, but sometimes 
serrate-crenate and with circular and linear pellucid dots 
scattered over the blade. The flowers are small and arranged 
in dense, axillary fascicles. The terminology relating to the 
structure of the flowers is not consistent, thus Phillips (l.c.) 
and Hutchinson (1967) refer to “calyx lobes”, Wild (l.c.) 
to “sepals” and Sleumer (l.c.) to “tepals”. Similarly 
Phillips, Hutchinson and Wild refer to “staminodes”, while 
Sleumer refers to “disc lacinia”. 

According to Palmer and Pitman (1973) the ashes of 
the bark of C. gladiiformis are used by the Bantu as snuff. 



Fig. 30. — 1, Casearia gladiiformis, flowering twig, x 2 /i (Moll 3290); a, flower, x 10 (Moll 3290); b, fruit, x 1 (Dutton 
22 ). 




by A. A. Obermeyer 

Herbs, shrubs or trees, usually pubescent. Leaves alternate, simple, variously incised or 
entire; stipules small or 0. Flowers regular, bisexual, in racemose or cymose, axillary or terminal 
inflorescences or solitary, opening in the morning, fading at night; pedicels free or rarely adnate to 
petiole; bracts small. Calyx tubular, 5-dentate or sepals 5, fused near the base, quincuncial. Petals 
5, free, erect or rotate, contorted in bud, usually unguiculate with the claw inserted on the 
calyx-tube or near its base, sometimes with a ligule or small thin corona on upper part of claw. 
Stamens 5, hypogynous or inserted on the calyx-tube, filaments free, of equal or unequal length, 
anthers bilocular, introrse, opening lengthwise, basifixed. Ovary superior, 1-locular, with 3 parietal 
or, very rarely, basal placentas; styles 3, usually free, slender, often heterostylous; stigmas 
multipartite; ovules many to few (-1). Capsule ovoid or silique-like (resembling Brassicaceae 
fruits), 3-valved, with 1- to multi-seriate placentas, dehiscent; seeds cylindrical, straight or curved, 
reticulate or striate, often pitted, on long funicles and enveloped by a white, leaf-like aril. 

Genera 9, species about 90, predominantly South American; also in Africa, the Mascarenes and south-east Asia. Four 
genera with 10 species recorded from the northern parts of South Africa and South West Africa. 

Literature: Urban in Jahrb. K. Bot. Gart. Mus. Berl. 2 (1883). 

Annuals or chamaephytes, 10-60 cm tall; flowers in axillary inflorescences or solitary; petals 
orange-red to pale yellow, claw not dark-coloured; pedicels free; 

Flowers on long racemes; capsule resembling a siliqua of the Brassicaceae, long, thin, 

terete, rostrate; seeds straight, cylindrical 1 . Wormskioldia 

Flowers solitary or on short, few-flowered racemes; capsule ovoid; seeds curved: 

Calyx tubular; petals not clawed; capsule erect, narrowly ovoid, beaked, thin-walled; 

leaves without 1-2 pairs of sessile glands below 2. Streptopetalum 

Calyx of 5 sepals fused in lower third; petals clawed, alternating with sepals; 
capsule recurved, ovoid, woody; leaves with 1-several pairs of round sessile glands 

below on lower half of leaf 3. Piriqueta 

Shrubs up to 2 m tall with axillary, solitary flowers; petals orange or yellow with a dark brown 

or purplish black claw; pedicels fused to petiole 4. Turnera 


Wormskioldia Thonn. in Schum. & Thonn., Beskr. Guin. PI. 165 (1827); Urb. in Jahrb. K. Bot. 
Gart. Mus. Berl. 2: 48 (1883); Gilg in Pflanzenfam. ed. 2, 21: 464 (1925); Burtt Davy, FI. Transv. 
1: 118 (1926); Phill., Gen. ed. 2: 517 (1951); Lewis in F.T.E.A. Turneraceae: 9 (1954); Hutch., 
Gen. Flow. PI. 2: 352 (1967). Type species: W. pilosa (Willd.) Schweinf. ex Urb. 

Annuals or perennials (often chamaephytes), variously hispid and often glandular-pubescent. 
Leaves alternate, linear to narrowly ovate, usually serrate or lobate to deeply incised; stipules small 
or 0. Racemes simple, secund, few- to many-flowered with the apical part recurved in bud, 
accrescent and erect with age, the flowers developing one after another, opening in the morning, 
fading at night; scape long and thin; pedicels short, somewhat accrescent and often recurved in 
fruit; bracts small; a few sterile bracts often present on the rhachis. Calyx tubular, 5-dentate, 
usually setiferous on the outside, pubescent in lower half inside. Corolla rotate, yellow, orange-red 
or white; petals obovate, unguiculate, the claw appressed to the calyx-tube in upper half, fused to it 



below; with a ligule or small pustule on upper part of claw. Stamens 5, hypogynous, reaching to the 
throat, filaments filiform, sometimes narrowly winged, occasionally 3 shorter; anthers oblong. 
Ovary ellipsoid, styles 3, erect, heterostylous, stigmas apical, small, multipartite. Capsule 
resembling a silique of Brassicaceae, terete, usually long and thin, often constricted between the 
uniseriate seeds, rostrate; seeds straight, terete, obtuse, reticulate, usually with 2 pores in each 
cavity, aril unilateral, short. 

An African genus with about 1 1 species in tropical and subtropical regions; recorded from the northern parts of South 
Africa and South West Africa. 


With bulbous-based setae on stems; exstipulate: 

Capsule erect, hairy; leaves coarsely serrate 1. W. glandulifera 

Capsule patent or reflexed, glabrous; leaves pinnatisect, discolorous, soft and thin 2. W. tanacetifolia 

Without bulbous-based setae on stems, hairs simple, short, sparse; leaves lobate in lower half, rarely 

entire; stipules auriculate, fused to base of leaf 3. W. lobata 

Perennials (chamaephytes) forming a woody basal complex and a long thick taproot; the parts above 
ground dying back annually; without bulbous-based setae: 

Leaves narrowly linear, sessile, entire, glabrous 4. W. mossambicensis 

Leaves broader, petiolate or sessile, margin variously incised, pubescent: 

Capsule c. 3 cm long, few-seeded, not constricted between the seeds; plants glandular-pubescent but 

without long setae; leaves coarsely and irregularly double-serrate 5. W. schinzii 

Capsule c. 5-8 cm long, many-seeded, constricted between the seeds; plants without glandular 
pubescence but with long dark red or white setae on the stems; leaves subentire to laciniate: 

Stems with long patent red to dark brown bristles, other parts glabrescent; leaves greyish-green, 

subentire to dentate, rarely laciniate; racemes 6-12-flowered 6. W. longipedunculata 

Stems, leaves, peduncles and capsules echinulate; bristles on stems not reddish; leaves 

yellow-green, laciniate; racemes 4— 7-flowered 7. W. lacerata 

1. Wormskioldia glandulifera Klotzsch 
in Peters, Reise Mossamb. Bot. 1: 146, t.26 
(1864); Mast, in F.T.A. 2: 503 (1871); Urb. in 
Jahrb. K. Bot. Gart. Mus. Berl. 2: 49 (1883); 
Gilg in Pflanzenfam. ed. 2, 21: 463 (1925); 
Burtt Davy, FI. Transv. 1: 1 19 (1926); A. & R. 
Fernandes in Mem. Junta Invest. Ultram. ser. 2, 
34: 16 (1962). Type: Mozambique, Tete, Peters 
(B, holo.f; K; EA; PRE, photo.). 

Small glandular-pubescent annuals up to 
30 cm tall, usually few-branched. Stems, 
petioles, veins, scapes and pedicels bearing 
erect multicelled setae which are swollen and 
dark basally, the capitate glands usually lost 
early on, interspersed with simple, soft hairs. 
Leaves petiolate, broadly linear-acuminate, 4—8 
cm long, 8-20 mm broad, coarsely and irregu- 
larly double-serrate, drying a dark brown col- 
our, lamina sparsely and softly puberulent. 
Flowers c. 3 on each axillary raceme; peduncle 
5-10 cm long, wiry, arcuate-erect; pedicels up 
to 5 mm in fruit; with minute bracts. Calyx 
tubular, c. 1 cm long, puberulent; lobes short, 
acute. Petals c. 1,5 cm long, yellow or orange. 
Stamens unequal. Ovary cylindrical, hairs 
erect; styles heterostylous. Capsule erect, over- 
topping the leaves, up to 6 cm long, puberulous; 
seeds narrowly clavate, cream, brown or black, 

the ridges minutely knobbed, aril tongue- 
shaped. Fig. 31: 1. 

Recorded from Mozambique, Rhodesia, South West 
Africa, Transvaal and Natal; behaving as a weed. 

S.W.A. — 1819 (Karakuwisa): Cigarette, N.E. of 
Karakuwisa, Maguire 2462. 1820 (Tarikora): Omuramba 
Khaudum at Tamso Camp, De Winter & Marais 4672. 
1918 (Grootfontein): near Tsebeb, Schoenfelder S536. 

Transvaal. — 2229 (Waterpoort): Dongola Hill, Codd 
& Dyer 3907; Liliput, Strey3486; Waterpoort , Bremekamp 
& Schweickerdt 297. 2230 (Messina): Messina, Rogers 
19405. 2231 (Pafuri): Pafuri, Van der Schijff3568; 5 km N. 
of Kloppersfontein, Acocks 16782. 2327 (Ellisras): 4 km 
N.E. of P.O. Tom Burke, Codd 6615. 2330 (Tzaneen): 
Hans Merensky Nature Reserve, Gilliland 780. 2331 
(Phalaborwa): Kruger National Park, near Letaba Camp, 
Munro s.n. 2530 (Lydenburg): 16 km N. of Abel Erasmus 
Pass on road to Tzaneen, Mauve 4328. 253 1 (Komatipoort): 
16 km from Gorge, Van der Schijff 2283. 

Natal.— 2731 (Louwsburg): Pongola Dam at Magut, 
Gerstner 3174. 

2. Wormskioldia tanacetifolia Klotzsch 
in Peters, Reise Mossamb. Bot. 1: 147 (1864); 
Urb. in Jahrb. K. Bot. Gart. Mus. Berl. 2: 51 
(1883); Burtt Davy, FI. Transv. 1: 119 (1926); 
A. & R. Fernandes in Mefn. Junta Invest. 
Ultram. 2, 21: 16 (1962). Type: Mozambique, 
between Boror and Querimba, Peters 23 (B, 
holo.f; K). 



Fig. 31. — Leaves and capsules of Wormskioldia spp., all x 1: 1 , W. glandulifera, leaf; la, capsule (Fan der Schijff 
1885); 2, W. tanacetifolia, leaf; 2a, capsule (Rogers 20830); 3, W. lobata, leaf; 3a, capsule (Welwitsch 2493); 4, W. 
mossambicensis, leaf; 4a, capsule (Vahrmeijer 1057); 5, W. schinzii, leaf; 5a, capsule ( Mostert 1239); 6, W. 
longipedunculata, leaf; 6a, capsule (Codd & Over 4698, Rogers 25503); 7, W. lacerata, leaf; 7a, capsule (Rogers 



Annuals up to 60 cm tall, malodorous. 
Stems, petioles, veins, scapes and pedicels bear- 
ing erect multicelled setae which are swollen 
and dark basally, the apical capitate glands 
usually early deciduous; with some soft, white 
woolly hairs on the stems. Leaves with petioles 
1-3 mm long; lamina somewhat ovate- 
acuminate in outline, c. 6 cm long and 4 cm 
broad, deeply bipinnatipartite, thin, soft, dis- 
colourous, nearly glabrous. Flowers in c. 
3-flowered patent racemes, longer than leaves, 
bracts small, circular. Calyx c. 16 mm long, 
glabrous. Petals yellow, c. 27 mm long. Sta- 
mens unequal with 2 longer than the others. 
Ovary glabrous. Capsule recurved, c. 3 cm 
long; seed oblong, brown or yellow. Fig. 31:2. 

Recorded from the eastern Transvaal, Mozambique 
and Rhodesia. Like W. glandulifera it behaves as a weed; 
often in sandy disturbed ground, emitting an unpleasant 

Transvaal. — 2530 (Lydenburg): Nelspruit, Liebenberg 
2465 ; Buitendag 402; Kaapse Hoop, Rogers 20830. 2531 
(Komatipoort): Storiwane, S.E. of Pretoriuskop, Van der 
Schijff 1927; 12 km W. of Malelane, Codd 5246; Louws 
Creek, Thorncroft 2044. 

3. Wormskioldia lobata Urb. in Jahrb. 
K. Bot. Gart. Mus. Berl. 2: 52 (1883); Thonner, 
Flow. PI. Afr. 1. 106 (1908); Lewis in F.T.E.A. 
Turneraceae: 10 (1954); A. & R. Fernandes in 
Mem. Junta Invest. Ultram. ser. 2, 28: 14 
(1961); ibid. 34: 15 (1962); Schreiber in 
F.S.W.A. 88: 4 (1968). Syntypes: several from 
tropical Africa including Angola, Pungo An- 
dongo, Welwitsch 2493, 2494 (Bf; BM; K; 

Annual up to 60 cm tall, minutely tawny 
pubescent with short and long hairs and with 
scattered small pyramidal, attenuate, multicellu- 
lar glands. Stems woody, few-branched. Leaves 
with the primary small, attenuate below, petiol- 
ate, exstipulate; following sessile, broadly 
linear, up to 12 cm long and 2 cm broad, apex 
acute, lower half usually expanded into 2 large 
unequal lobes, occasionally with a few shallow 
indentations above, margin shallowly serrate, 
soft, discolourous; stipules small, circular, 
fused to lamina below. Flowers 2-6 on long 
thin scapes overtopping the leaves. Calyx tubu- 
lar, c. 15 mm long, lobes short, acute, setose. 
Petals orange (or pale yellow in specimens 
from S.W.A.) up to 2 cm long. Stamens nar- 
rowly winged. Ovary glabrous. Capsule patent 
to reflexed, up to 8 cm long, glabrous. Fig. 31: 

Recorded from South West Africa. Widespread in 
tropical Africa, in sandy habitats. 

S.W.A. — 1718 (Kuring-kuru): near Mupini Mission, De 
Winter & Marais 5433; Musese Camp W. of Lupala 
Mission, De Winter & Marais 4975. 1721 (Mbambi); 64 
km W. of Andara, Merxmiiller & Giess 2074. 1724 
(Katima Mulilo): Ngoma area, Killick & Leistner 3005. 
Grid uncertain: Kaokoveld, Kunene River bank, Barnard 

4. Wormskioldia mossambicensis A. & 

R. Fernandes in Bol. Soc. Brot. Ser. 2, 35: 158, 
t.5 (1961); in Mem. Junta Invest. Ultramar, ser. 
2, 34: 18 (1962). Type: Mozambique, Inham- 
bane, between Quissico and Chiducuane, Men- 
donca 3308 (LISC, holo.). 

Chainaephyte, slender, fastigiate, glauc- 
ous, glabrous to slightly woolly pubescent. 
Stems c. 40 cm tall, thin, hard, striate. Leaves 
narrowly to broadly linear, variable, 8-15 cm 
long, 2-8 mm broad, margin entire or shallowly 
denticulate (the teeth consisting of glands), 
sessile; stipules minute, pectinate, sparsely la- 
nate inside axil. Flowers on long erect scapes, 
2-7-flowered. Calyx c. 1 cm long with a few 
short ovoid-attenuate glands, lobes with hyaline 
minutely fimbriate margins. Petals orange, c. 
12 mm long. Stamens equal or unequal. Ovary 
glabrous. Capsule c. 7 cm long, reflexed. Fig. 

Recorded from Mozambique and Natal in Palm veld; 
grassy sandy localities. 

Natal. — 2632 (Bela Vista): Tongaland, north of Lake 
Sibayi, Vahrmeijer & Tolken 274; 5 km W. of Maputa, 
Strey & Moll 3788. 2732 (Ubombo): S. of Maputa on road 
to Lake Sibayi, Gerstner 4114; Manzengwenya, Moll 
4717; Bazwana, Gerstner 3557; 707; Strey 5115. 2831 
(Nkandla): Hlabisa Bird Sanctuary, Gerstner 5027. 2832 
(Mtubatuba): W. of Charters Creek, Ward 2832; 26 km 
N.E. of Mtubatuba, A cocks 13085; 21 km N. of bridge over 
St. Lucia estuary on road to Cape Vidal, Codd 10168. 

5. Wormskioldia schinzii Urb. in Bot. 
Jahrb. 15: 159 (1892); in Warb., Kunene-Samb. 
Exped. 310 (1903); A. & R. Fernandes in Mem. 
Junta Invest. Ultramar, ser. 2, 28: 15 (1961). 
Type: Angola, Gambos, Calculovar River, 
Newton 26 (Z, holo.!; COI; PRE, photo.); 
incorrectly stated by Urban as collected in 

W. juttae Din ter & Urb. in Feddes Repert. 13: 153 
(1914); Schreiber in F.S.W.A. 88: 4 (1968). Syntypes: S.W. 
Africa, between Neitsas and Otjituo, Dinter 652 (Bf; 
SAM!), 652a (Bf). W. rehmii Suesseng. in Mitt. Bot. 
Staatssamml. Miinchen 1: 55 (1950). Type: S.W. Africa, 
farm Rotenfels, Rehm s.n. (M, holo.). 



Small scabrid and glandular-pubescent, 
spreading chamaephytes c. 30 cm tall. Stems 
densely leafy, woody and brittle. Leaves linear, 
9-18 cm long (rarely shorter), 12-25 mm 
broad, apex acuminate, base acute, margin 
coarsely double-serrate, the teeth diminishing in 
size towards the apex, scabrid, midrib raised, 
often winged below; petiole short or 0; stipules 
0. Flowers in few-flowered racemes. Calyx 16 
mm long. Petals c. 35 mm long, bright orange 
red. Ovary densely covered with erect bristles. 
Capsule c. 3-4 cm long, few-seeded, not con- 
stricted between seeds, glandular-pubescent. 
Fig. 31:5. 

Recorded from South West Africa, Botswana, northern 
Cape, Transvaal and Angola. In sandy soil. 

S.W.A. — 1718 (Kuring-kuru) or 1818 (Tsitsib): on road 
from Tsintsabis to Kuring-kuru, Schoenf elder S528. 1917 
(Tsumeb): Farm Kameeldoom, GR 158 , Seydel 2146. 1918 
(Grootfontein): Neitsas, Dinter 652. 2017 (Waterberg): 
near Okakarara, Liebenberg 4730. 2119 (Epukiro): Farm 
Hekel, G0415, Giess 10667. 2218 (Gobabis): Gobabis 
district, farm Gemsbokfontein, Merxmuller & Giess 1180. 

Transvaal. — 2326 (Mahalapye): Buffelsdrift, Vahr- 
meijer 1297. 

Cape. — 2523 (Pomfret): Bray, Mostert 1239. 

6 . Wormskioldia longipedunculata 

Mast, in F.T.A. 2: 502 (1871), as longepedun- 
culata; Urb. in Jahrb. K. Bot. Gart. Mus. Berl. 
2: 53 (1883); Burtt Davy, FI. Transv. 1: 119 
(1926) excl. f. 11a; Lewis in F.T.E.A. Tur- 
neraceae: 14 (1954); A. & R. Fernandes in 
Mem. Junta Invest. Ultram. ser. 2, 34: 18 
(1962); Letty, Wild Flow. Transv. 224, t.lll 
(1962). Type: Malawi, Manganja Hills, Meller 
(K, holo.; PRE, photo.). 

Small chamaephytes up to 40 cm tall, 
covered with a short white woolly pubescence 
and scattered short to long hairs; upper part of 
stem armed with c. 3 mm long dark red to 
purple, coarse patent hairs. Leaves close to- 
gether, sessile, variable, linear to oblong, 8-15 
cm long, 2-20 mm broad, apex usually obtuse, 
margin subentire to serrate to coarsely pinnat- 
ifid in lower half, glaucous, minutely white 
woolly pubescent and with some short scattered 
hairs, becoming glabrescent with age. Raceme 
7-15-flowered. Calyx-tube 15 mm long, 
setulose, Petals up to 24 mm long, bright 
orange-red. Stamens unequal in length. Ovary 
with a few scattered setae, probably heterostyl- 
ous. Capsule 5-8 cm long, indistinctly 
moniliform, many-seeded. Fig. 31: 6. 

Recorded from South West Africa, the Transvaal, 
Rhodesia, Malawi, Mozambique, Zambia and Botswana, in 
dry bushveld, road verges and lands. 

S.W.A. — 1723 (Singalamwe): On the road from Sibinda 
to Katima Mulilo, Pienaar & Vahrmeijer 224. 1724 
(Katima Mulilo): about 8 km from Katimo Mulilo to 
Liny anti, Killick & Leistner 3 101 . 

Transvaal. — 2228 (Maasstroom): road from Steilloop 
Bridge to Tolwe, Ihlenfeldt 2118a. 2230 (Messina): Sibasa, 
Junod 4547; Makonde, Codd 6871. 2231 (Pafuri): Punda 
Milia, Lang sub TRV 32178; Schlieben 9283. 2328 (Balti- 
more): Blouberg, Schlechter 4645. 2329 (Pietersburg): 
Lejuma Mt., Strey 7991 ; Sand River near Mara, Schlieben 
& Strey 8290; Soutpansberg, 8 km N. of Louis Trichardt, 
Van Vuuren 1664; Spelonken (Cave Mts.), Nelson 122; 
near Pietersburg, Bolus 10895. 2330 (Tzaneen): Elim, 
Obermeyer sub TRV 29324; Tshakhuma, Van Warmelo 
5337/11; Mbayinbayi, Lang sub TRV 32254; Mokeetsi, 
Van Dam sub TRV 25021; Westfalia Estate near 
Duiwelskloof, Scheepers 24; 13 km S.W. of Leydsdorp, 
Codd & Dyer 4698. 2428 (Nylstroom): Moorddrif, Leen- 
dertz sub PRE 7316. 2429 (Zebediela): Naboomfontein, 
Galpin 133156; 13 km N.E. of Zebediela, Story 5355. 2430 
(Pilgrims Rest): Shilouvane, Junod 4220; Marouvuni, 
Junod 85. 

Because of its soboliferous habit and deep hard woody 
roots it is able to persist in ploughed lands and on road 
verges, being considered a pest by farmers and attractive to 
passersby who admire the bright red flowers. It is said to be 
poisonous to cattle but the unpleasant odour acts as a 

7. Wormskioldia lacerata Oberm. in 
Bothalia 11: 288 (1974). Type: Transvaal, 
Newington, Rogers 22578 (PRE, holo.; NH). 

W. schinzii sensu Burtt Davy, FI. Transv. 1: 

1 19 (1926) inch f. 1 la. 

Chamaephytes with a dense short echinu- 
late pubescence, c. 30 cm tall. Stems usually 
short, firm, erect or ascending. Leaves close 
together, sessile, linear-lacerate, 8-12 cm long, 
2-3,5 cm broad, the lobes patent, irregular, 
dentate, decreasing in size upwards, apex 
acuminate, echinulate along the broad midrib, 
veins and margin, with a white woolly pubes- 
cence on the midrib and lateral veins above. 
Flowers 4—7 on long straight or slightly arcuate 
scapes, ultimately exceeding the leaves; bracts 
small. Calyx c. 15 mm long, with erect setae 
and some sparse white woolly pubescence. 
Petals deep orange, the blade broadly obovate, 
c. 2 cm long. Stamens 8 mm long in short- 
styled flowers, 4 mm long in long-styled flow- 
ers. Ovary densely pubescent with erect hairs, 
heterostylous, the styles 9 mm long and ex- 
serted above the stamens or 4 mm long, reach- 
ing only about half way. Capsule reflexed, c. 7 
cm long, echinulate. Fig. 31:7. 

Recorded from the eastern Transvaal and Swaziland, in 
open spaces in subtropical woodland or in disturbed places. 



Transvaal. — 2330 (Tzaneen): Hans Merensky Nature 
Reserve, Oates 77; 5 km S. of Gravelotte, Leach 11559; 
Werdermann & Oberdieck 1922. 2430 (Pilgrims Rest): 10 
km from Mica on road to Hoedspruit, Ihlenfeldt 2303; 16 
km from Mariepskop on road to Klaserie, Coetzer 151. 
2431 (Acomhoek): Klaserie, Killick & Strey 2511; 
Newington, Rogers 22578; Buitendag 909; Skukuza, 
Cholmondeley s.n. 2531 (Komatipoort): Pretoriuskop, Van 

der Schijff 1075; 1217; Codd <6 De Winter 4918 ; Malelane, 
Lang sub TRV 31644; Kaapmuiden, Rogers 25034; De 
Kaap Valley, Thorncroft 54; Barberton, Galpin 747; 
Thorncroft sub Rogers 19171 . 

Swaziland. — 2531 (Komatipoort): Tshaneni, near Sand 
River reservoir, Edwards 213. 2631 (Mbabane): Ranches 
Ltd., Compton 24704. 


Streptopetalum Hochst. in Flora 24: 665 (1841); Urb. in Jahrb. K. Bot. Gart. Mus. Berl. 2: 55 
(1883); Gilg in Pflanzenfam. ed. 2, 21: 463 (1925); Phill., Gen. ed. 2: 517 (1951); Lewis in 
F.T.E.A. Turneraceae: 15 (1954); Hutch., Gen. Flow. PI. 2: 352 (1967). Type species: S. serratum 

Small glandular-pubescent annuals or perennials. Stems woody. Leaves alternate, exstipulate. 
Flowers in few- to many-flowered axillary racemes. Calyx narrowly tubular, hairy on the outside 
and within in lower half. Corolla rotate, petals 5, convolute in bud, inserted in the throat of the 
calyx, spathulate, yellow or orange. Stamens 5, free, perigynous, equal, or unequal in length, 
reaching to the mouth of the tube. Ovary oblong-ovoid, pubescent, with 3 parietal placentas; 
ovules pluriseriate; styles filiform just over-topping the stamens. Capsule ellipsoid, 3-valved, 
splitting to the base, glandular-hispid; seeds allantoid, testa reticulate, pits with 2 pores, aril 
unilateral, as long as, or shorter than the seed, with a wavy margin. 

Species 3, African; 1 species recorded from Ethiopia to the Transvaal and South West Africa. 

Urban and others observed 5 large tubercles on the calyx-tube above the insertion of the stamens but in the 
(unfortunately) few flowers dissected, these appeared to be absent. 

Streptopetalum serratum Hochst. in 
Flora 24: 666 (1841); Urb. in Jahrb. K. Bot. 
Gart. Mus. Berl. 2: 56 (1883); Lewis in 
F.T.E.A. Turneraceae: 17 (1954); Schreiber in 
F.S.W.A. 88: 2 (1968). Type: Ethiopia, Tsel- 
lenti, Takkaze River, Schimper 1260 (B, 
holo.f; K; BM). 

Small bushes up to 40 cm tall, annual or 
possibly sometimes perennial, sparsely to den- 
sely hispid with short simple hairs and glan- 
duliferous setae with swollen bases, many- 
celled. Leaves sessile, narrowly ovate, up to 10 
cm long and 2,5 cm broad, attenuate below and 
above, serrate, thin, yellow-green. Inflores- 
cences with the primary racemes axillary, sim- 
ple, several-flowered, secund; the inner serial 
axillary bud later develops into a strong side 
branch which in its turn produces much ab- 
breviated side branches bearing reduced leaves 
and a few flowers towards the top. Calyx 
narrowly tubular, c. 1 cm long, lobes 2,5 mm 

long, hairy outside and inside below. Petals 
spathulate, c. 7 mm long, yellow or orange, 
alternating with sepals. Stamens 5, attached just 
below the middle of the tube, free above, 
reaching to the throat. Ovary ovoid, densely 
covered with glandular setae which are swollen 
at the base; styles 3, terete, reaching to the 
throat; stigmas apical, obtuse, slightly swollen. 
Capsule ovoid, c. 12 mm long, rostrate, scabrid; 
seeds allantoid, 2,5 mm. Fig. 32. 

Recorded from northern and central Transvaal, Bot- 
swana, South West Africa and further north to Ethiopia; rare 
but often locally abundant, in warm dry bushveld, usually in 
sandy soil, often along road sides. 

S.W.A. — 1917 (Tsumeb): on sandy road from Tsumeb to 
Heidelberg, Dinter 7530; Tsumeb, Dinter 3034, 1714. 

Transvaal. — 2228 (Maasstroom): Kremetartberg, 

Mara Research Station 475. 2230 (Messina): Messina, 
Rogers 20820. 2231 (Pafuri): 16 km E. by N. of Punda 
Milia, Acocks 16775. 2329 (Pietersburg): Blouberg, Mogg 
24548. 2428 (Nylstroom): Magabene, Schlechter 4671 . 
2529 (Witbank): Rooikop, Smuts & Gillett 2536. 

5357 3. PIRIQUETA 

PiriquetazlM^/., Hist. PI. Guiane Fr. 1: 298, 1. 117 (1775); Urb. in Jahrb. K. Bot. Gart. Mus. Berl. 
2: 57 (1883); Gilg in Pflanzenfam. ed. 2, 21: 463 (1925); Phill., Gen. ed. 2: 517 (1951); A. & R. 
Fernandes in Mem. Junta Invest. Ultram. ser. 2, 34: 22 (1962). Type species: P. villosa Aubl. 



Fig. 32.— 1, Streptopetalum serratum, habit, X 1; a, longitudinal section showing attachment of filaments, x 6; b. ovary 
and styles, x 4; c, capsule, x 3; d, seed with aril, X 14 (Mara Research Station 475). 



Erblichia Seem., Bot. Voy. Herald 130 t.27 (1853) pro minore parte quoad species austro-africanae; Hutch., Gen. Flow. 
PI. 2: 353 (1967). 

Annuals, shrubs or trees, variously pubescent with simple and/or compound hairs. Leaves 
alternate, sessile or petiolate, stipules minute or 0, lamina simple, serrate, with or without large 
sessile circular glands on lower surface. Flowers axillary, solitary or rarely cymose. Calyx forming 
a short to long tube. Petals inserted in the throat, obovate, clawed, alternating with the calyx-lobes 
and together forming a fimbriate annulus in the throat. Stamens hypogynous with long or short 
filaments. Ovary with 3 parietal placentas, ovules many, uni- to multi-seriate; styles 3, 
heterostylous. Capsule globose, 3-valved, hairy; seed straight or slightly curved, reticulate, aril 
unilateral, as long as or shorter than seed. 

Species about 20, mostly South American. One species in Transvaal, Natal, Swaziland and 

Hutchinson places Piriqueta capensis in the genus Erblichia in his Gen. Flow. PL 2: 353 (1967), but without 
validating the new combination. Urban and Gilg regard Erblichia as a section of Piriqueta. Four species are involved; the 
type species, E. odorata from Central America and 2 indigenous in Madagascar, all 3 of which fall outside the scope of this 
work. Our species, the fourth, forms a short but distinct calyx-tube and a fimbriate annulus in the throat, characters which 
Urban considers of importance for Piriqueta. Harvey overlooked this annulus for it is small and unobtrusive. The 
differences enumerated by Hutchinson in some cases do not agree with those observed by Urban. There appears to be no 
reason why it should not be placed under Piriqueta. 

Piriqueta capensis (Harv.) Urb. in Jahrb. 
K. Bot. Gart. Mus. Berl. 2: 78 (1883); Gilg in 
Nat. Pflanzenfam. ed. 2, 21: 464 (1925); Fer- 
nandes in Bol. Soc. Brot. Ser. 2, 35: 160, t.7 
(1961); in Mem. Junta Invest. Ultram. ser. 2, 
34: 22 (1962). Type: Transvaal, Apies River, 
Burke & Zeyher (TCD, holo.! K, SAM!, PRE, 

Tumera capensis Harv. in F.C. 2: 599 (1862); Thes. Cap. 
2: 25, 1. 140 (1862). 

Small woody, sericeous, aromatic shrub- 
lets (chamaephytes) 15-30 cm high, with a long 
thick woody taproot and gnarled basal stems 
covered with thick corky bark, producing an- 
nual shoots. Leaves subsessile, stipules reduced 
to 1-2 pairs of minute, subulate glands, lamina 
narrowly ovate, 2-5 cm long, 7-14 mm broad, 
apex acute to obtuse, base cuneate, margin 
serrate, with 1-2 pairs of sessile circular glands 
irregularly placed on lower surface near margin, 
densely pubescent with coarse simple hairs. 
Flowers axillary, solitary, opening in the morn- 
ing, fading at night, on short to long pedicels 
which become pendulous in fruit, with a pair of 
minute bracts. Sepals fused in lower third 
forming a cup, free above, narrowly ovate- 
acuminate, apiculate, c. 12 mm long, 3 mm 
broad, thin, dorsally somewhat hairy, 3-nerved, 
deciduous. Petals about as long as sepals, 
yellow, narrowly obovate, clawed, fused to the 

sepaloid cup below, alternating with the sepals 
and together producing a small erect, annular 
fringe at the mouth of the cup, deciduous. 
Stamens 5, free, hypogynous, filaments sub- 
ulate, glabrous, long or short, anthers versatile, 
introrse. Ovary ovoid, densely covered with 
erect hairs; ovules 4-8, more or less uniseriate 
on each parietal placenta; styles 3, long or short 
(heterostylous), terete; stigmas spreading- 
fimbriate. Capsule ovoid, 8 mm, the valves 
splitting to the base; seeds oblong often some- 
what curved, 3 mm, yellow or black, reticulate, 
with a short areole. 

Recorded from the Transvaal, Swaziland, northern 
Natal, Mozambique and Rhodesia, usually on stony hill- 
sides in dry bushveld. Fig. 33. 

Transvaal. — 2229 (Waterpoort): between Waterpoort 
and Wylliespoort, Obermeyer , Schweickerdt & Verdoorn 
339. 2327 (Ellisras): farm Alfred, 4 km W. of Monte 
Christo P.O., Codd 6609; near Elmeston, about 64 km from 
Vaalwater, Meeuse & Strery 10432. 2330 (Tzaneen): Sand- 
rivier, Schlechter 4597. 2428 (Nylstroom): between 

Warrribad and Nylstroom, Bremekamp & Schweickerdt 3; 
Side y 1312; Bolus 11890. 2429 (Zebediela): 9 km N.W. of 
Marble Hall, Codd & Verdoorn 10371. 2528 (Pretoria): 7 
km N.E. of Hammanskraal on road to Rust de Winter, Codd 
3619. 2529 (Witbank): Loskop Dam on road to “The 
Hell”, Codd & Verdoorn 10361. 

Swaziland. — 2631 (Mbabane): Palata near Stegi, Comp- 
ton 31209. 

Natal. — 2732 (Ubombo): Ingwavuma hills, Strey 



Fig. 33. — 1, Piriqueta capensis, habit, x %; a, leaf-base, 
seed with aril, x 13 ( Strey 10263). 

lower surface showing glands, X 2; b, flower laid out, x 3; c. 



5360 4. TURNERA 

Turnera L., Sp. PI. 1: 271 (1753); Benth. & Hook.f., Gen. PI. 1: 806 (1862); Urb. in Jahrb. K. 
Bot. Gart. Mus. Berl. 2: 81 (1883); Gilg in Pflanzenfam. ed. 2, 21: 464 (1925); Hutch., Gen. Flow. 
PI. 2: 352 (1967). Type species: T. ulmifolia L. 

Annual or perennial herbs, shrubs or trees, pubescent. Leaves alternate, sessile or petiolate, 
entire, toothed or subpinnatifid, often with 2 basal glands on the lower side at the base, pubescence 
simple or stellate; stipules small or 0. Flowers axillary, solitary, rarely forming inflorescences, the 
peduncle sometimes fused to the petiole, bracteate. Calyx forming a short tube, 5-partite. Petals 5, 
inserted in the calyx-throat, shortly clawed. Stamens inserted below the petals, sometimes 
hypogynous. Ovary ovoid to cylindrical, ovules pluriseriate; styles 3, spreading; stigmas fimbriate. 
Capsule globose to ovoid, seeds globose to obovoid or cylindrical and curved, reticulate, arillate. 

Species about 100, mostly South American; one species endemic in South West Africa and Angola, another in tropical 
Africa. One South American species naturalized in the Seychelles, Bourbon and Indo-Malaysia. 

Turnera oculata Story in Bothalia 7: 493 
(1961). Type: S.W. Africa, Kaokoveld, Kunene 
River at Otjinungua, De Winter & Leistner 
5770 (PRE, holo.; WIND). 

Shrub up to 2 m tall, with slender long 
erect branches and short side branches, covered 
by a silvery (turning tawny when dry) velvet 
pubescence of stellate and simple sericeous 
hairs or hair-covering more sparse; intemodes 
often shortened above. Leaves obovate, up to 3 
cm long and c. 12 mm broad, apex obtuse to 
acute, base narrowed into a petiole, margin 
shallowly to deeply dentate, velvety pubescent 
to sparsely pubescent with short stellate hairs 
and long simple sericeous hairs, pale green, 
nerves distinct below, with 2 (rarely 1 or 0) 
conspicuous sessile circular, rimmed glands, 1 
mm in diam. at junction of petiole and blade; 
stipules minute. Flowers solitary, axillary on 
the upper abbreviated side branchlets; pedicels 
fused to petioles; bracts 2, filiform, c. 10-18 
mm long. Calyx tubular, c. 12 mm long, 9 mm 
in diam., 10-nerved, lobes linear-acuminate, 
somewhat longer than tube, pubescent with a 
membranous glabrous margin. Petals convo- 
lute, inserted on the calyx throat, spreading 
above, broadly obovate, c. 2,5 cm long and 1,8 
cm broad, apex obtuse, apiculate, base cuneate, 
bright yellow above, dark red-brown below 
forming the dark heart of the flower. Stamens 5, 
exserted, filaments 3 cm long, arcuate, inserted 
3 mm below throat with the shortly winged 
margins adhering to the calyx-tube below, an- 
thers basifixed, the filaments inserted in a dorsal 
pit, introrse. Ovary cylindric, obtuse, pubes- 
cent, ovules pluriseriate, numerous; styles 3, 
longer than stamens, spreading; stigmas 
fimbriate. Capsule 3-valved, ovoid, c. 9 mm 
long, pubescent and pustulate; seed cylindrical, 

somewhat curved, 4 mm long, cream, reticu- 
late, aril unilateral, membranous, lobed, longer 
than seed. 

Recorded from the Kaokoveld in northern South West 
Africa and from southern Angola. Two varieties are recog- 

Pubescence velvety, dense; leaf margin with shallow 

teeth; bracts c. 10 mm long (a) var. oculata 

Pubescence more sparse; leaf margin with distinct 
sharp teeth; bracts 10-19 mm 
long (b) var. paucipilosa 

(a) var. oculata. 

Turnera oculata Story in Bothalia 7: 493 (1961); Fer- 
nandes in Mem. Junta Invest. Ultram. ser. 2, 34: 22 (1962). 
Schreiber in F.S.W.A. 88: 3 (1968). Type: S.W. Africa, 
Kaokoveld, Kunene River at Otjinungua, De Winter & 
Leistner 5770 (PRE, holo.; WIND). 

Plants with a densely velvety pubescence 
hiding the epidermis; teeth on leaf-margin small 
and shallow; bracts up to 10 mm long. Fig. 34: 
1 . 

Recorded from the Kaokoveld in northern South West 
Africa and southern Angola. 

S.W. A. — 1712 (Posto Velho): 1 km S. of Kunene River 
on the mountain E. of Otjinungua, Giess & Wiss 3290; De 
Winter .& Leistner 5770 ; Otjihipa Mountain, Davies, 
Thompson & Miller 77. 1812 (Sanitatas): 20 km S. of 
Kunene River on way to Anabib (Orupembe), Story 5778. 

(b) var. paucipilosa Oberm. in Bothalia 
11: 290 (1974). Type: S.W. Africa, near Ot- 
jitanda, Giess 9377 (PRE, holo.; WIND). 

Plants sparsely and minutely pubescent 
with stellate and simple hairs not hiding the 
epidermis; teeth on leaf margin usually deeply 
incised; bracts 12-18 mm long. Fig. 34: 2. 

Recorded from one area in the Kaokoveld, around 
Etengua and Otjitanda. 

S.W. A. — 1712 (Posta Velho): Otjinungua, 12 km S. of 
Kunene River, De Winter <5 Leistner 5751. 1713 

(Swartbooisdrift): 32 km W. of Etengua near Otjitanda, 
Giess 9377; 8 km W. of Otjitanda, Kotze 118. 




Fig. 34. — 1, Turnera oculata var. oculata, branch with flower, x V4; la, base of leaf, lower surface showing glands, X 4; 
lb, anther, X 10; lc, capsule and bracteoles, x lVi; Id, capsule and leaf showing fusion of petiole and pedicel, x 2; le, 
seed with aril, X 10 ( Story 5778). 2, T. oculata var. paucipilosa, leaf, x 1; 2a, capsule and bracteoles, x 1 (Giess 




by W. J. J. O. De Wilde* 

Climbers or sometimes (erect) herbs, mostly provided with axillary tendrils (tribe Pas- 
sifloreae), or erect shrubs or trees (tribe Paropsieae); glabrous or hairy, rarely thorny. Leaves 
(mostly) alternate, simple or compound, entire or lobed, often with glands on petiole and blade; 
stipules small, sometimes caducous. Inflorescences axillary, either cymose (Passifloreae), ending 
in (a) tendril(s) or not, or racemose (Paropsieae); bracts and bracteoles usually small. Flowers 
hermaphrodite or (functionally) unisexual (and then plants often dioecious or polygamous); stipe 
articulate to pedicel; hypanthium saucer-shaped to tubiform. Sepals 4 or 5 (or 6), imbricate, free or 
partially connate ( Adenia p.p.), often persistent. Petals (3-4 or 5 (-6), imbricate, rarely absent. 
Corona extra-staminal, inserted on the hypanthium, various, composed of hairs, or of one or more 
whorls of thread-like processes or scales, or tubiform or cup-shaped, or absent {Adenia p.p.). Disc 
mostly extra-staminal, annular or composed of 5 mostly strap-shaped parts {Adenia), or absent. 
Stamens 4— many, in S. Africa 4 or 5 (-8), inserted on the hypanthium or on an androgynophore, if 
few altemipetalous, free or partially connate; anthers 2-celled, basifixed to dorsifixed, versatile or 
not, sometimes apiculate, opening lengthwise. Ovary superior, sessile or on a gynophore or 
androgynophore, 1 -celled, 3-5 (-6)-carpellate, with 3-5 (-6) parietal placentas; ovules mostly 
numerous, anatropous, with 2 integuments; styles 1 or 3-5, very short to distinct, free or partially 
united; stigma(s) capitate to subglobose, sometimes much divided {Adenia). Fruit a loculicidally 
3-5-valved capsule, or berry-like. Seeds mostly compressed, enveloped by a membranous or 
mostly pulpy aril; funicles often distinct, testa crustaceous, mostly pitted; endosperm homy; 
embryo large, straight, with foliaceous cotyledons. 

A pantropical family of about 18 genera and about 500 species; Passiflora, the largest, mainly in America. 

The family can be divided into two tribes, viz. tribe Paropsieae, and tribe Passifloreae. Tribe Paropsieae (all African, 1 
species of Paropsia in the Caprivi-area) comprises the erect, arborescent species, destitute of tendrils. Tribe Passifloreae 
consists of the mainly climbing species, provided with tendrils; of this tribe 3 genera are indigenous, whereas Passiflora is 
cultivated as an ornamental or for the flavoured edible fruit (P. edulis Sims), but sometimes escaped and running wild. Two 
species (P. foetida L. and P. suberosa L.) are locally naturalized, and may behave as weeds. 

Shrubs or trees, without tendrils; vegetative ramification through the axillary bud; corona 

tomentose, composed of threads arranged in 5 bundles (Tribe Paropsieae) ... .1. Paropsia 
Mostly tendril climbers, rarely low herbs without tendrils, or shrub-like, sometimes provided 
with thorns; vegetative ramification through the accessory bud (Tribe Passifloreae): 

Flowers (mostly) unisexual; corona ill-developed, or absent; disc consisting of 5 strap- 
shaped parts, or absent; stigmas divided or distinctly papillate 2. Adenia 

Flowers hermaphrodite; corona well-developed; disc annular or absent; stigmas capitate or 

Styles single, stigma 3 or 4-lobed 3. Schlechterina 

Styles 3 or 4 (or 5), free or partially connate: 

Flowers small, up to 1,5 cm diam.; androgynophore absent, stamens inserted inside in 

the inner corona 4. Basananthe 

Flowers usually much larger; androgynophore distinct 5. Passiflora 

*Rijksherbarium, Schelpenkade 6, Leiden, Netherlands. 



5369a 1. PAROPSIA 

Paropsia Noronha exThouars, Hist. Veg. Isles Austr. Afr., ed. 1, 59, tab. 19 (1805); ed. 2, 59, 
tab. 19 (1806); Gilg in Pflanzenfam. ed. 2, 21: 414 (1925); A. & R. Fernandes in Garcia de Orta 6: 
244, 667 (1958); Sleumer in Bull. Jard. Bot. Nat. Belg. 40: 50 (1970). Type: P. edulis Thouars. 

Trichodia Griff, in Not. 4: 570 (1854). 

Hounea Baill. in Bull. Soc. Linn. Paris 1: 301 (1881). 

Shrubs or trees, mostly hairy, without tendrils. Leaves alternate, ± distichous, not lobed, ± 
oblong, petiolate, margin subentire to mostly distinctly serrate-dentate or repand-dentate; glands at 
the tops of the teeth, and on the blade margin (rarely on the surface) towards the base; stipules 
small, mostly caducous. Inflorescences 1-many-flowered, subfasciculate, axillary, peduncled or 
not, yellow to reddish-brown hairy, sometimes forming terminal panicles by reduction or 
abscission of sustaining leaves; bracts linear, caducous. Flowers hermaphrodite, pedicellate, often 
fragrant; hypanthium shallowly cup-shaped; sepals (4 or) 5, imbricate, persistent; petals (4 or) 5, 
imbricate. Corona composed of hairy threads in one series, ± free or connate at base, or these 
collected into 5 bundles opposite the petals. Stamens 5, opposite the sepals; filaments glabrous or 
pilose at base, faintly connate at base, inserted at base of ovary or on short gynophore; anthers 
oblong to linear, ± sagittate, subdorsifixed. Gynophore short or absent; ovary ovoid, placentas 
3-5, many-ovuled; styles (2 or) 3(-5), free, sometimes pilose; stigmas subreniform or capitate. 
Fruit a 3(-5)-valved capsule, subglobose, sometimes shortly stipitate. Seeds ovoid, compressed, 
5-7 mm, arillate; testa crustaceous, scrobiculate. 

A genus of 1 1 species, 1 in S.E. Asia (Malesia), 6 species in Madagascar, and 4 in tropical Africa, 1 of which enters our 
Flora area.* 

Paropsia brazzeana Baill. in Bull. Soc. 
Linn, Paris 1: 611 (1886); Warb. in Pflanzen- 
fam. 3, 6a: 27 (1893); Gilg in Bot. Jahrb. 40: 
472 (1908); in Engl., Pflanzenw. Afr. 3, 2: 572, 
fig. 253 D-F (1921); Gilg in Pflanzenfam. ed. 2, 
21: 415 (1925); Exell in J. Bot. Lond. 67 
(Suppl.): 191 (1929); Gossw. & Mendonca, 
Carta Fitogeogr. Angola 111, 161 (1939); 
Gossw., Agron. Angol. 7: 274 (1953); A. & R. 
Fernandes in Garcia de Orta 6: 653 (1958) as 
brazzaeana\ F. White, For. FI. N. Rhod. 268 
(1962); Sleumer in Bull. Jard. Bot. Nat. Belg. 
40: 70 (1970). Type: Congo Republic, De 
Brazza 48 (P, holo. !). 

P. reticulata Engl, in Bot. Jahrb. 4: 391 (1892); ibid. 15: 
586 (1893); Warb. Pflanzenfam. 3, 6a: 27, fig. 9D-F 
(1893); Gilg. in Warb., Kunene-Samb. Exped. 309(1903); 
Pflanzenfam. ed. 2, 21: 414, fig. 183D-F (1925). Type: 
Angola, Mechow 541 (B+; K, photo.). — var. ovatifolia 
Engl, in Bot. Jahrb. 14: 14 (1891). Type: Zaire, Pogge 951 
(Bf). — var. proschii Briq. in Annu. Cons. Jard. Bot. 
Geneve 6: 1 (1902). Type: Zambia, Prosch 35 (G). P. 

*After going to press specimen Wiens & Van Wyk 
5334 from Pafuri (Grid 2231 AD) was received. The 
specimen is immature and has been provisionally identified 
as P. brazzeana Baill. by the author. In the editor’s opinion the 
specimen probably belongs to P. braunii Gilg, a species 
known from Mozambique and areas further north. 

argutidens Sleumer in Fedde Repert. 45: 13 (1938). Type: 
Zambia, Milne-Redhead 726 (K, holo. ! ; BR). 

Shrub or shrublet 1-2,5 (-3) m; stem up to 
5 cm across, mostly branched from the base; 
young twigs yellowish-tomentose. Leaves 
oblong-elliptic or ovate-oblong, rarely broadly 
elliptic or ovate, apex shortly acuminate, rarely 
obtuse, base broadly attenuate to rotundate, 
hard-chartaceous, when young woolly 
yellowish-tomentose all over, later on glabres- 
cent and shiny above, margin regularly glandu- 
lar sinuate-dentate or subserrate (teeth 1-4 mm, 
2-5 (-7) mm spaced), (3-) 6-10 (-13) x (1,5-) 
2,5-4 (-6,5) cm (in sterile shoots somewhat 
larger), lateral nerves 5-8 pairs, upward curved, 
conjunct, distinctly prominent beneath; petioles 
(3-) 5-7 mm long, c. 2 mm thick. Inflores- 
cences 1-2 (-3) (rarely-5)-fiowered fascicles, 
axillary to normal leaves; bracts ovate, c. 2 mm. 
Flowers subsessile or stipe (pedicel) up to 5 
mm, c. 1 mm thick; sepals oblong, greenish- 
whitish, outside (as the pedicel) yellowish- 
tomentose, inside finely tomentose, 10-12 x 
(3-) 4-5 mm, slightly enlarging with age; petals 
oblong, white, especially outside puberulous, 
resembling the sepals but slightly narrower; 
corona c. 3 mm high, split up into 5 rather 



distinct bundles; threads connate at base, out- 
side tomentose, connate part inside glabrous; 
filaments c. 5 mm; anthers oblong-cordate, c. 2 
mm; ovary sessile, reddish-brown tomentose; 
styles hardly 1 mm. Fruit ovoid-subglobose, 
red-brown tomentose, 1,5-2 x 1,2-1, 8 cm. 
Fig. 35. 

Distributed mainly in Equatorial Africa, viz. Came- 
roon, Central African Republic, Congo Brazzaville, Zaire, 
Angola, Zambia, Rhodesia, Botswana; 1 collection from 
Caprivi; found in forest and thickets, gallery- and secondary 
forest; rather common on clay and Kalahari sand; 0-1000 
(in Angola up to 1600) m. 

S.W.A. — 1723 (Singalamwe): Caprivi area, Singalamwe 
to Katima Mulilo, Killick & Leistner 3281. 

5370 2. ADENIA 

Adenia Forsk., FI. Aegypt.-Arab. 77 (1775); Engl, in Bot. Jahrb. 14: 374 (1891); Harms in 
Pflanzenfam. ed. 2, 21: 488 (1925); Burtt Davy, FI. Transv. 1: 220 (1926); Liebenberg in Bothalia 
3: 513 (1939); A. & R. Fernandes in Garcia de Orta 6: 255, 656 (1958); C.F.A. 4: 217 (1970); 
Phill. , Gen. ed. 2: 518 (1951); De Wilde in Meded. Landbouwhogeschool Wageningen 71-18: 
1—28 1 (1971); Ross, FI. Natal 251 (1972). Type: A. venenata Forsk. 

Modecca Lam., Encycl. 4: 208 (1797); Harv. in F.C. 2: 499 (1862); Thes. Cap. 2: 43 (1863); Gen. PI. ed. 2: 121 (1868); 
Benth. & Hook.f., Gen. PI. 1: 813 (1867); Mast, in F.T.A. 2: 512 (1871). 

Kolbia P. Beauv., FI. d’Oware 2: 91, 1. 120 (1807). 

Blepharanthes J. E. Smith, Gram. Bot. 188 (1821), nom. illegit. 

Paschanthus Burch., Trav. 1: 543 (1822). 

Microblepharis (Wight & Am.) Roem., Syn. Mon., 2 Pepon. 200 (1846). 

Erythrocarpus Roem., Syn. Mon., 2 Pepon. 204 (1846). 

Clemanthus Klotzsch in Peters, Reise Mossamb. Bot. 143 (1862). 

Ophiocaulon Hook.f. in Benth. & Hook.f., Gen. PI. 1: 813 (1867); Harv., Gen. PI. ed. 2: 121 (1868); Mast, in F.T.A. 2: 
517 (1871); Engl, in Bot. Jahrb. 14: 385 (1891). 

Machadoa Welw. ex Benth. & Hook.f., Gen. PI. 1: 814 (1867). 

Keramanthus Hook.f. in Bot. Mag. t.627 1 (1876). 

Jaggia Schinz in Verh. Bot. Ver. Prov. Brandenb. 30: 253 (1888). 

Echinothamnus Engl, in Bot. Jahrb. 14: 383 (1891). 

(Sub)ligneous to herbaceous perennial climbers with tendrils, sometimes erect herbs or 
shrublets mostly without tendrils, often with a rootstock or tuber, or a pachypodous main stem; 
sometimes thorny or spiny; glabrous or sometimes pubescent. Leaves either simple, entire or 
lobed, or palmately parted or pseudo-compound; glands (0 or 1) or 2 at the blade-base, at or near 
the apex of the petiole, and with or without glands elsewhere on the lower surface or margin of the 
blade; stipules minute, narrowly triangular or reniform; tendrils axillary. Inflorescences axillary, 
cymose, the middle (or the first three) flower(s) often replaced by (a) tendril(s); bracts and 
bracteoles minute, triangular to subulate. Flowers dioecious or rarely monoecious, bisexual or 
polygamous, campanulate or urceolate to tubular or infundibuliform, mostly greenish or yellowish, 
always glabrous, stipe articulate at base. Hypanthium saucer- or cup-shaped, or tubular. Sepals 
(4—) 5 (-6), free or partially connate into a calyx-tube, imbricate, persistent. Petals (4—) 5 (-6), 
free, included in the calyx, sometimes adnate with the calyx-tube, mostly fimbriate or laciniate. 
Corona annular, or consisting of 5 cup-shaped parts, or of a laciniate rim or membrane, or 
composed of hair-like processes, or absent. Disc-glands 5, ligulate or strap-shaped, inserted at or 
near the base of the hypanthium, alternating with the petals, or absent. Male flowers: Stamens (4-) 
5 (-6), hypogynous or perigynous (variably inserted in the hypanthium), free or partially connate 
into a tube; anthers basifixed, oblong to linear, often apiculate, 2-locular, opening introrsely to 
latrorsely; vestigial ovary minute. Female flowers mostly smaller than the male flowers, with 
smaller petals; staminodes ± subulate. Ovary superior, shortly stipitate or subsessile, globose to 
oblong; placentas 3 (-5), ovules usually numerous; styles 3(-5) free or partially united, sometimes 
styles very short; stigmas mostly subglobose, laciniate or plumose or densely woolly-papillate. 



Fig. 35. — 1, Paropsia brazzeana, flowering twig, x 1; a, flower, x 2% (Flanagan 3205); b, fruit, x 1; c, seed, x 3 
(Jacobsen 1726). 



Fruit a stipitate 3 (-5)-valved capsule; pericarp (woody-)coriaceous to rather fleshy (and hence 
fruit ± berry-like), greenish to yellow or bright red. Seeds ± compressed, with crustaceous pitted 
testa, enclosed in a membranous to pulpy (juicy) aril. 

A genus of 93 species, divided into 6 sections, in tropical Africa (with a few species in temperate Southern Africa), 
Madagascar, and S.E. Asia, Malesia, and N. Australia. 10 species occur in our area. 

Gland at blade-base single, on a single median hemispherical to spathulate appendage; corona absent; disc-glands 

absent; sepals free; stigmas (sub)sessile. Sect. Ophiocaulon 10. A. gummifera 

Gland(s) at blade-base (0 or) 1 or 2, (sub)sessile, never on a single distinct median appendage with narrow insertion; 
corona absent or membranous or laciniate, or composed of (fine) hairs; disc-glands present or not; sepals free or 
partially connate into a calyx-tube; styles distinct: 

Sepals free or nearly so; sepals and petals inserted at or at about the same level; anthers wholly or largely extending 
beyond the hypanthium; corona present; disc-glands present. Sect. Microblepharis: 

Male flowers including stipe usually more than 10 mm long; plants mostly provided with tendrils, or with thorns: 
Plant unarmed: 

Gland at blade-base 1; leaves entire or 3-5-foliolate; hypanthium wider than long, 2,5-4 mm 

wide 1. A. fruncosa 

Glands at blade-base 2, though approximate; leaves deeply 5-lobed; hypanthium longer than or about as long 

as wide, 2-3 mm wide 2. A. glauca 

Plant thorny 3. A. spinosa 

Male flowers small, including stipe 8-10 (-14) mm long; stems without tendrils, erect, short, hard-shrubby, 

arising from a swollen lumpy base; glands at blade-base 2, sessile 4. A. pechuelii 

Sepals partially connate into a calyx-tube which extends wholly or partially above the insertion of the petals, hence 
calyx-lobes and petals not inserted at the same level, but if approximate because of adnation of the petals to the 
calyx-tube, then stamens (anthers) largely or entirely enclosed in the calyx-tube: 

Midrib not ending in a gland; stamens inserted at the base of the hypanthium; corona present; disc-glands present. 

Sect. Blepharanthes: 

Leaves (deeply) lobed or entire, not digitately compound: 

Hypanthium wider than long, 5-12 mm wide, base flattish, about as wide as the calyx-tube; leaves 

3-5-lobed 5. A. natalensis 

Hypanthium about as long as, or longer than wide, 3-6 mm wide, base tapering; leaves entire . .6. A. hastata 
Leaf-blade digitately dissected to the base, the lobes or leaflets sometimes more or less stalked: 

Climbing herbs 1-5 m tall, provided with tendrils; leaves palmately (3-) 5-parted or -foliolate . .7. A. digitata 

Erect herbs, up to 50 cm tall, without tendrils; leaves palmately (5-) 7-parted 8. A. wilmsii 

Midrib ending in a (sub)apical gland; stamens inserted well above the base in the flower tube; corona absent; 
disc-glands absent. Sect. Paschanthus 9. A. repanda 

1. Adenia fruticosa Burn Davy, FI. 
Transv. 1: 36; 221 (1926); Bremekamp, Vege- 
tationsbilder 23, 3: 6, pi. 18 (1932); Lieben- 
berg in Bothalia 3: 538, 528, 532, pi. 1, 2 
(1939); Letty, Wild Flow. Transv. 225 (1962); 
De Wilde in Meded. Landbouwhogeschool 
Wageningen 71-18: 69 (1971). Type: Trans- 
vaal, Pole Evans H15723 (K, holo.!; PRE!). 

Shrub or shrubby tree to 6 m, glabrous; 
main stem mostly thick, soft-woody, branched 
or not, up to 2 x 0,6 m; branches shrubby or 
lianoid, provided with tendrils. Leaves some 
grey-green or glaucous, punctate or not be- 
neath, simple or 3-5-foliolate, in outline ovate 
to suborbicular, base (sub)cordate, 1-8 x 1-8 
cm, (1-) 3-5-plinerved; petiole (0,3-) 1-5 cm; 
leaflets (or lobes) orbicular, ovate or obovate, 
base rounded to attenuate, apex subacute to 
broadly rounded, rarely retuse, 1-6 x 1-4 (-6) 

cm, ± penninerved; margin entire; petiolule 
0-5 (-7) mm. Gland at blade-base 1, on a 
fleshy ± upward curved median lobelet at the 
top of the petiole; no other glands present. 
Stipules narrowly triangular, c. 0,5 mm. 
Inflorescences either solitary in the leaf-axils, or 
grouped in short-shoots, peduncled up to c. 1 
cm, 2-5-flowered in 6, 1-3-flowered in ?; 
tendrils 0; sterile tendrils 3-12 cm, sometimes 
breaking off and leaving a thorn-like structure. 
Male flowers campanulate, including the 0,5-5 
mm long stipe 9-17 mm; hypanthium cup- 
shaped, 1—2,5 x 2, 5-4, 5 mm; calyx-tube 0; 
sepals lanceolate, obtusish, 7-9 (-10) mm, 
(sub)entire; petals oblong to lanceolate, acute, 
4—8 mm, 3-nerved, serrulate; filaments 1,5-4 
mm, connate for 0,5-2 mm, inserted at the base 
of the hypanthium; anthers 3-6,5 mm, obtuse; 
septa 1-2 mm high; corona consisting of fine 
hairs 0,2-0, 5 (-1) mm, sometimes partly mem- 



branous; disc-glands c. 0,5 mm, or absent. 
Female flowers campanulate, including the 
0,5-1 mm long stipe 7-9 mm; hypanthium 
1-1,5 x 2, 5-3, 5 mm; calyx-tube 0; sepals 
lanceolate, obtuse, 5-7 mm, entire; petals lan- 
ceolate, acute, 3-4 mm, 1 -nerved, ± serrulate; 
staminodes 2-4 mm; septa 1 (-2) mm high; 
corona consisting of fine hairs 0,2-0, 5 mm, or 
nearly absent; disc-glands 0-0,5 mm; 
gynophore 1,5-3 mm; ovary subglobose 2,5-4 
mm; styles connate for 0,5-0, 7 mm, style arms 
0,5-1 mm; stigmas subglobose, papillate, each 
1-1,5 mm diam. Fruit 1-2 per inflorescence, 
subglobose (to ellipsoid), excluding the 2-5 
mm long gynophore 1-2 x 0,8-1, 8 cm; 
pericarp coriaceous; seeds 3-6 per capsule, 
broad ovate to orbicular, 6-6,5 mm. 

A species of dry savanna and bushveld, on sandy or 
granitic soils; 100-1400 m. Distributed in Transvaal and 
Natal, also in Rhodesia. 

Three more or less allopatric subspecies are recog- 

Leaves 3-5-foliolate; petiolule of leaflets 2-5 (-7) mm; 

anthers c. 3 mm (a) subsp. fruticosa 

Leaves simple or 3 (or 4)-foliolate; leaflets ses- 
sile; anthers 4-5,5 mm: 

Hypanthium of 3 flowers broadly cup-shaped, ± 
5-saccate; corona hairs 0,5-1 mm; disc-glands 
present; leaves simple or 3- 

foliolate (b) subsp. simplicifolia 

Hypanthium of 3 flowers cup-shaped, taper- 
ing, not saccate: corona hairs up to 0,5 mm, or 
partly absent; disc-glands absent; leaves 3 (or 
4)-foliolate (c) subsp. trifoliolata 

(a) subsp. fruticosa. 

De Wilde in Meded. Landbouwhogeschool 
Wageningen 71-18: 70 (1971). 

Leaves 3-5-foliolate, 2-8 x (2-) 2,5-8 
cm; petiole 1-5 cm; leaflets suborbicular to 
obovate, base attenuate to rounded, apex 
broadly rounded, rarely retuse, 1-6 x 1-4 (-6) 
cm; petiolules 2-5 (-7) mm. Male flowers 
including the c. 0,5 mm long stipe c. 9 mm; 
hypanthium cup-shaped, not distinctly saccate, 
c. 1 x 2,5-3 mm; sepals 7-8 mm; petals c. 4 
mm; filaments c. 1,5 mm; anthers c. 3 mm; 
corona partly consisting of fine hairs up to 0,5 
mm, partly membranous; disc-glands c. 0,2 
mm. Female flowers including the 0,5-1 mm 
long stipe 7-9 mm, hypanthium c. 1,5 x 

2. 5- 3, 5 mm; sepals 5-7 mm; petals 3-4 mm; 
staminodes 3—4 mm; corona consisting of fine 
hairs up to 0,2 mm; disc-glands absent; pistil 

6.5- 8 mm; gynophore 2-3 mm; ovary 3-4 x 
3-3,5 mm; styles connate for 0,7 mm, style- 
arms 0,5-0, 7 mm. 

Found in bushveld, on dry stony slopes, dry sandy 
loam soil and granite in the eastern Transvaal, not 
elsewhere; 800-1400 m. Flowers in Aug. and Sept., fruits in 
Oct. and Nov. 

Transvaal. — 2429 (Zebediela): Chuniespoort, Ober- 
meyer & Verdoorn 10; Sekukuniland, Barnard 454. 2430 
(Pilgrims Rest): 8 km N. of Burgersfort, Codd & Dyer 
7732; Kasparsnek, Strey 3693. 

(b) subsp. simplicifolia De Wilde in 
Meded. Landbouwhogeschool Wageningen 
71-18: 71 (1971). Type: Rhodesia, Chase 1321 
(SRGH, holo. !; BM; K; LISC). 

Leaves simple, ovate, or (2-) 3-foliolate, 
1-6 x 0,7-5 cm; petiole (0,3-) 0,5-2, 5 cm; 
leaflets suborbicular to elliptic, or ovate, or 
rhomboid, base attenuate to rounded, apex 
(sub)obtuse, 1-5 x 1-3 (-4) cm; petiolules ± 
0. Male flowers including the 3-4 mm long 
stipe 13-15 mm; hypanthium broadly cup- 
shaped, 5-saccate, c. 2 x 4-4,5 mm; sepals 
8-10 mm; petals 7-8 mm; filaments 3-4 mm; 
anthers 5-5,5 mm; corona of densely set fine 
hairs 0,5-1 mm; disc-glands 0,5-0, 7 mm. 
Female flowers including the 0,5-1 mm long 
stipe 7-8 mm; hypanthium 1-1,5 x 2, 5-3,5 
mm; sepals 5-6 mm; petals 3-4 mm; 
staminodes 2-2,5 mm; corona hairs c. 0,5 mm; 
disc-glands 0, 2-0,5 mm; pistil 5-6,5 mm; 
gynophore 1,5-2 mm; ovary 1,5-3 x 1,5-2, 5 
mm; styles connate for 0,5-0, 7 mm; style-arms 
0,5-1 mm. 

Confined to a restricted area in Transvaal and southern 
Rhodesia, in thorny bushveld, found in rocky (basalt) places 
and on sandy soils, also near hot springs; 400-1000 m. 
Flowers and fruits Sept.-Nov. 

Transvaal. — 2229 (Waterpoort): Dongola Reserve, 
Pole Evans 3747. 

The fruits are reported as being green berries whitish 

(c) subsp. trifoliolata De Wilde in 
Meded. Landbouwhogeschool Wageningen 
71-18: 72 (1971). Type: Natal, Acocks 12937 
(PRE, holo.!; P). 

Leaves 3- (or 4)-foliolate, 1,5-3 X 1,5-2, 5 
cm; petiole 0,5- 1,5 cm; leaflets suborbicular, 
base and apex rounded to subacute, 0,5-2 x 
0,5-2, 5 cm; petiolules ± 0. Male flowers in- 
cluding the c. 5 mm long stipe c. 14—17 mm; 
hypanthium cup-shaped, tapering, not saccate, 
c. 2,5 x 4—5 mm; sepals 6,5-8 mm; petals c. 5 
mm, filaments 3-3,5 mm; anthers 4-4,5 mm; 
corona hairs up to 0,5 mm, or corona partly 
membranous or partly absent; disc-glands 0. 
Female flowers not known. Fruit subsessile, 
ovoid, c. 1,2 cm long, 2-3-seeded. 



An endemic of Natal and Zululand, in sandy bushveld 
on sandstone; locally frequent; 100-900 m. Flowers and 
fruits Sept.-Dee. 

Natal. — 2732 (Ubombo): Ubombo Mt., Venter 1751. 
2831 (Nkandla): Umfolozi Game Reserve, Ward 4378; near 
Ntonjaneni, A cocks 12937. 2832 (Mtubatuba): Hluhluwe 
Game Reserve, Ward 4475. 

Fresh fruits are recorded as: “almost globose, mottled 
green when immature, darker towards base, becoming 
orange-green when ripe”, or: “very dark green with light 
green veins”. 

Gerstner 5241 is annotated as having an underground 
tuber, and not an overground stem. 

2. Adenia glauca Schinz in Bot. Jahrb. 
15, Beibl. 33, 1; 1 (1892); Engl., Pflanzenw. 
Afr. 3, 2: 605 (1921); Burtt Davy, FI. Transv. 1: 
222 (1926); Steyn, Tox. PI. S. Afr. 314, fig. 
43B, 46, 47B (1934); Liebenberg in Bothalial: 
539, 523, 532, fig. ^8, pi. 4 & 5 (1939); Letty, 
Wild Flow. Transv. 225, tab. 112 (1962); Watt 
& Breyer-Brandwijk, Med. Pois. PI. ed. 2: 828 
(1962); De Wilde in Meded. Landbouwhoge- 
school Wageningen 71-18: 73 (1971). Type: 
Transvaal, Rehmann 4799 (Z, lecto. !). 

Modecca glauca Schinz in Bot. Jahrb. 15, Beibl. 33, 1: 1 
(1892); ex Engl, in Bot. Jahrb. 14: 393 (1891), nom. nud. 

Climber, sometimes shrub-like, up to 3,5 
m, the basal part thickened, tapering, “bottle- 
shaped”, up to c. lx 0,4 m, ± fleshy, smooth, 
greenish, glabrous. Leaves greyish or glaucous 
or purplish-grey, sometimes punctate beneath, 
5-parted to the base, suborbicular in outline, 

2- 12 x 2,5-12 cm, base cordate, 5-plinerved; 
petiole (0,5-) 1-5 cm; leaf-parts (ob)ovate to 
oblong, ± attenuate at base, apex (sub)obtuse, 
sometimes retuse, 1-7 x 0,5-3, 5 (-4,5) cm; 
margin entire. Glands at blade-base 2, contigu- 
ous, situated on two ± connate thickish auricles 
making the blade slightly peltate; blade-glands 
1-3 (-5), (sub)apical on the lobes. Stipules 
narrowly triangular, acute, 1-1,5 mm. Inflores- 
cences peduncled up to 1 (-2) cm, 2-5-flowered 
in S, 1-3-flowered in ?; tendril (0 or) 1, 2-6 cm; 
sterile tendrils up to 10 cm. Male flowers 
(tubiform to) infundibuliform, including the 

3- 7 mm long stipe (13-) 15-30 mm; hypan- 
thium cup-shaped, (1-) 2-4 x 2-4 mm; calyx- 
tube 0; sepals lanceolate, obtuse, (8-) 10-18 
(-20) mm, entire; petals lanceolate, apex acute 
to obtuse, 6-11 mm, 1-3-nerved, serrulate; 
filaments (3-) 4,5-7 mm, connate for (1,5-) 

2,5-4 mm, inserted at the base of the hypan- 
thium; anthers 3 — 5,5 mm subacute; septa (1-) 

2,5-4 mm high; corona consisting of either 

sparse thick hairs c. 0,5 mm or of thinner hairs 
0,5-1 mm; disc-glands absent. Female flowers 
tubiform to infundibuliform, including the 

1.5- 4 mm long stipe (9-) 10-15 mm; hypan- 
thium cup-shaped, 1,5-2 x 2-3 mm; calyx-tube 
0; sepals lanceolate, obtuse, 7-9 mm, (sub)en- 
tire; petals lanceolate, acute, 4—5 mm, 1-nerved, 
(sub)serrulate; staminodes 2-3,5 mm; septa 1-2 
mm high; corona hairs sparse, thick, 0,5-1 mm; 
disc-glands 0; gynophore 1-2 mm; ovary sub- 
globose, 2, 5-4, 5 mm; styles connate up to 0,5 
mm, style-arms 1-1,5 mm; stigmas subglobose, 
papillate, each c. 1,5 mm diam. Fruit 1-2 per 
inflorescence, (subglobose to) ellipsoid, exclud- 
ing the 3-5 mm long gynophore 1,8—2, 5 X 

1.5- 2 cm; pericarp coriaceous; seeds 3-5 per 
capsule, orbicular to heart-shaped, 5, 5-7, 5 mm. 

Found in a restricted area in Transvaal and S.E. 
Botswana, in dry bushveld, in rocky places or on sandy soil; 
locally common; 1 000-1 600 m. Flowers mainly Aug.- 
Jan., fruits Oct. and Nov. 

Transvaal. — 2427 (Thabazimbi): 17 km S. of Mat- 
labas, Codd 4433. 2428 (Nylstroom): near Naboomspruit, 
Galpin 11605; near Warmbaths, Burtt Davy 2622; Acocks 
13923. 2528 (Pretoria): Premier Mine, Rogers 25027; 
Magaliesberg, C. A. Smith 6271. 

Said to be poisonous to catde. 

Liebenberg (l.c.) mentions occasional 4-merous 
ovaries and fruits, and once a male flower with 6 petals and 

3. Adenia spinosa Burtt Davy, FI. 
Transv. 1, 36: 222 (1926); Liebenberg in 
Bothalia 3: 533, 523, fig. 9, pi. 3 (1939); Letty, 
Wild Flow. Transv. 225 (1962); De Wilde in 
Meded. Landbouwhogeschool Wageningen 
71-18: 76 (1971). Type: Transvaal, Rogers 
19299 (K, holo. !; PRE; Z). 

Thorny shrub up to 1,5 (-6?) m high, 
arising from an irregularly shaped ± fleshy 
trunk up to 0,5 m tall, up to 2,5 m wide. Leaves 
± coriaceous, grey-glaucous, sometimes 
punctate beneath, entire, broadly ovate to ellip- 
tic, base cordate to rounded, apex obtuse, some- 
times retuse, 1-3,5 x 0,7-3 cm, 3-plinerved to 
± penninerved; margin entire; petiole 0, 2-0,7 
cm. Gland(s) at blade-base 1 or 2, sessile at the 
top of the petiole; blade-gland 1, (sub)apical. 
Stipules triangular, acute, c. 0,5 mm. Thorns 
0,5-4 cm, patent, acute, when young ending in 
a tendril. Inflorescences solitary or in short- 
shoots, peduncled up to 0,5 cm, 2-6-flowered in 
6, 1-3-flowered in ?; tendril (0 or) 1, strong, 

4-6 cm, mostly replaced by a thorn; sterile 
tendrils simple, 4—8 cm. Male flowers 
tubiform-campanulate, including the 1,5-3 mm 


long stipe (10-) 12-24 mm; hypanthium (nar- 
rowly) cup-shaped, 1-3 X 2,5-4 mm; calyx- 
tube 0; sepals lanceolate, obtuse, (8-) 9-18 
mm, (sub)entire; petals lanceolate, acute, (6-) 
8-10 mm, 1-3-nerved, serrulate; filaments 

2, 5-5, 5 mm, connate for 1-3 mm, inserted at or 
near the base of the hypanthium; anthers 4—5 
mm, obtuse; septa 1-3 mm high; corona con- 
sisting of a few stiff hairs 0,2-1 mm, mainly 
near the insertion of the petals; disc-glands 
absent or inconspicuous. Female flowers 
tubiform-campanulate, including the 1-1,5 mm 
long stipe 7-10 mm, hypanthium 1-1,5 mm, 
calyx-tube 0; sepals lanceolate, obtuse, 5-8 
mm, entire; petals lanceolate, acute, 2, 5-3, 5 
mm, 1-nerved, (sub)entire; staminodes c. 1,5 
mm; septa 0,5-0, 7 mm; corona absent or con- 
sisting of a few stiff hairs c. 0,2 mm; disc- 
glands 0; gynophore 1-2 mm; ovary sub- 
globose, 2-4 (-5) mm; styles connate for c. 1 
mm, style-arms c. 0,5 mm; stigmas subglobose, 
papillate, each c. 1 mm diam. Fruit 1-2 per 
inflorescence, subglobose (to ellipsoid), exclud- 
ing the 3-4 mm long gynophore 1,4— 2,2 x 
1,2—1, 8 cm; pericarp coriaceous; seeds 3-6 per 
capsule, suborbicular, c. 6,5-7 mm. 

Occurs in a restricted area in the Transvaal, also in 
southern Rhodesia. It is locally common in open bushland 
and scrub, in dry rocky (basalt) places; 200-1000 m. 
Flowers in Jan., May, July, Sept, and Nov., fruits in Sept, 
and Nov. 

The thorns and tendrils are homologous with the 
inflorescences as found in other Adenias. 

The flowers resemble those of A. glauca. The mostly 
early deciduous leaves often much resemble the simple 
leaves of A. fruticosa subsp. simplicifolia. 

Transvaal. — 2220 (Waterpoort): Soutpan, Obermeyer, 
Schweickerdt & Verdoorn 137; Galpin 15140; 6 km N.W. 
of Wylliespoort, Codd 2997. 2230 (Messina): near Messi- 
na, Rogers 19299; 21664. 2329 (Pietersburg): Naauwpoort, 
Bremekamp & Schweickerdt 469; Vivo, Mogg 24453 . 2331 
(Phalaborwa): Kruger National Park, near The Gorge Rest 
Camp, Codd 6187. 2429 (Zebediela): Molsgat, near 
Chuniespoort, Obermeyer sub TRV 34667. 

4. Adenia pechuelii (Engl.) Harms in 
Pflanzenfam. 3, 6a, Nachtr. 1; 255 (1897); ibid., 
ed. 2, 21; 490, fig. 216 (1925); Engl., Pflan- 
zenw. Afr. 3, 2: 595, 600, 601, 608, fig. 267 
(1921); Schreiber in F.S.W.A. 89: 2 (1968); De 
Wilde in Meded. Landbouwhogeschool 
Wageningen 71-18: 78 (1971). Syntypes: S.W. 
Africa, Pechuel-Loesche s.n.; Giirich 18 (Bf). 

Echinotharnnus pechuelii Engl, in Bot. Jahrb. 14: 383, 
fig. 9 (1891). 

1 1 1 

Bushy shrub up to 1,5 m, mostly leafless, 
without tendrils; pruinose, glabrous; the 
branches up to 60 cm, thorny, growing from 
lumpy subglobose to cylindrical main stem up 
to 1 m diam. Leaves coriaceous, grey-green, 
minutely papillate beneath, entire or shallowly 
3-lobed, ovate-oblong to lanceolate, base at- 
tenuate to rounded, apex (sub)acute to obtuse, 
1-6 x 0,2-2 (-2,5) cm, penninerved; margin 
entire; petiole up to 0,2 cm. Glands at blade- 
base 2, lateral at the very base of the midrib; 
blade-glands 0-4, one pair at about V 3 -V 2 , and 
one pair 2 / 3 from the base. Stipules narrowly 
triangular, acute, 0,5-1 mm. Inflorescences 
peduncled up to 0,2 cm, 1-2 (-3)-flowered in 6 
and $; tendrils 0. Male flowers campanulate, 
including the c. 3 mm long stipe 8-10 (-14) 
mm; hypanthium cup-shaped, 0,7-2 x 2,5-4 
(-6) mm; calyx-tube 0; sepals elliptic to oblong, 
obtuse, 3-5,5 (-7) mm, entire; petals lanceol- 
ate, acute, 3-5 mm, 1-nerved, subserrulate; 
filaments 1-2 mm, connate for 0,5-1 mm, 
inserted at the base of the hypanthium; anthers 

2. 5- 4, 5 mm, obtuse; septa 0,5-1 mm high; 
corona hairs fine, 0,5-1 mm; disc-glands c. 0,2 
mm. Female flowers campanulate, including 
the 2-3 mm long stipe 7-8 mm; hypanthium 
(broadly) cup-shaped, 1-1,5 x 2,5-3 mm; 
calyx-tube 0-0,5 mm, sepals oblong, obtuse, c. 
4 mm, entire; petals lanceolate, (sub)acute, c. 
2,5 mm, 1-nerved, ± serrulate; staminodes 

1.5- 2 mm; septa 0,5-0, 7 mm high; corona hairs 
c. 0,5 mm; disc-glands 0; gynophore c. 1 mm, 
ovary subglobose to ellipsoid, c. 2 mm; styles 
connate for c. 0,5 mm, style-arms c. 0,5 mm; 
stigmas subglobose, papillate, c. 1 mm diam. 
Fruit 1 per inflorescence, subglobose to ellip- 
soid, excluding the 1-2 mm long gynophore 
1-1,7 x 1-1,5 cm; pericarp thinly coriaceous; 
seeds 3-10 per capsule, ovate-subcircular, 5-7 

A local endemic species in South West Africa, in 
semi-desert on sand, marble, and quartz and granite rock; 
200-1000 m. Flowers in Dec., fruits in Febr. 

S.W. A. — 2115 (Karibib): Omaruru, Giess 9190; Pforte, 
Merxmiiller 1758. 2214 (Swakopmund): near Walvis Bay, 
Jensen s.n. 2215 (Trekkopje): near Husab, Pechuel- 
Loesche s.n. 2416 (Maltahohe): Blasskranz, Volk 931. 

5. Adenia natalensis De Wilde in 
Meded. Landbouwhogeschool Wageningen 
71-18: 161, fig. 25 (1971). Type: Natal, Ger- 
rard 1820 (K, holo. !; BM). 

Subherbaceous climber, c. 4 m; glabrous. 
Leaves grey-green, not punctate beneath. 



deeply 3-5-palmately lobed or -partite, orbicu- 
lar to 5-angular in outline, base subcordate, 5-9 
x 6-10 cm, 5-subplinerved; lobes or blade- 
parts ovate to (ob)lanceolate, entire or 2-4 
(-6)-lobed, apex rounded to subacute, 2-6 cm; 
margin entire; petiole 1,5-6 cm. Glands at 
blade-base 2, separate or ± contiguous, on the 

1- 2 mm wide peltate blade-base; blade-glands 

2- 6, submarginal or ± scattered. Stipules sub- 
triangular, c. 0,5 (-1) mm, withering. Inflores- 
cences peduncled for 1-4,5 cm, (1-) 2-6- 
flowered in <3; tendril 0 or 1, 2-5 cm, sterile 
tendrils 8-12 cm. Male flowers campanulate, 
including the 4-4,5 mm long stipe 20-22 mm; 
hypanthium saucer-shaped, ± 5-saccate, 2-3 x 
10 (-13) mm; calyx-tube (1,5-) 2-3 mm; 
calyx-lobes elongate-triangular, (sub)acute, 
10-12 mm, subentire; petals obovate-oblong or 
± spathulate-unguiculate, apex rounded, 7,5-9 
mm, 3-5-nerved, denticulate, inserted at the 
same level as the corona; filaments 4—5 mm, 
connate for 1,5-2 mm, inserted at the base of 
the hypanthium; anthers c. 9 mm, (sub)obtuse; 
septa 1,5-2 mm high; corona consisting of hairs 
simple or branched, or of a laciniate membrane, 
(0,5-) 1 mm; disc-glands 1,7-2 mm. Female 
flowers and fruit not known. 

This species is known from only two collections made 
in Natal, in the previous century; apparently it is a local 
endemic and rare species, not unlikely extinct by now. 
Presumably it is a species of low coastal forest. 

Natal. — Zululand: without precise locality, Gerrard 
1200; 1820. 

6. Adenia hastata (Harv.) Schinz in Bot. 
Jahrb. 15, Beibl. 33, 1: 3 (1892); Burtt Davy in 
Ann. Transv. Mus. 3: 121 (1912); Engl., Pflan- 
zenw. Afr. 3, 2: 603 (1921); Harms in Notizbl. 
Bot. Gart. Berl. 8: 295 (1923); in Pflanzenfam. 
ed. 2, 21: 491 (1925); Burtt Davy, FI. Transv. 1: 
221 (1926); Liebenberg in Bothalia 3: 536, fig. 
4—5 (1939); A. & R. Fernandes in Garcia de 
Orta 6: 256 (1958); Watt & Breyer-Brandwijk, 
Med. Pois. PI. ed. 2: 828 (1962); De Wilde in 
Meded. Landbouwhogeschool Wageningen 
71-18: 168 (1971). Type: Natal, Gerrard 1199 
(TCD, holo.; BM; K; W). 

Modecca hastata Harv., Thes. Cap. 2: 43, tab. 167 

Adenia schlechteri Harms in Bot. Jahrb. 33, 1: 150 
(1902). Type: Transvaal, Schlechter 11747 (BM; BR; 
HBG; K; P; Z). 

Herbaceous climber to 4 m, growing from 
a tuberous rootstock; glabrous. Leaves pale- or 

greyish-green, often punctate beneath, simple, 
broadly ovate to ± hastate, base cordate to 
truncate, apex obtuse to acute, up to 2 mm 
subapical mucronate, 1,5-10 (-14) x 1,5-10 
(-13) cm, 3-7-(sub)plinerved; margin entire; 
petiole (0,5-5 (-10) cm. Gland(s) at blade-base 
1 or 2, contiguous, situated on the not- or 
shortly-peltate, sometimes 2-lobed blade-base; 
blade-glands 0-2 (-4), often conspicuous, 
(sub)marginal or near the top; leaves of juvenile 
specimens with peltate base, without glands. 
Stipules narrowly triangular, acute, 1,5-2 mm. 
Inflorescences peduncled for 0,5-4 (-11) cm, 
up to 12-flowered in S, 1-3-flowered in ?; 
tendril (0 or) 1, 1-5 cm; sterile tendrils up to 12 
cm. Male flowers tubiform to infundibuliform, 
including the 2-6 mm long stipe (13-) 15-30 
cm; hypanthium cup-shaped, tapering, 2-3 mm; 
calyx-tube (5-) 7-15 x (3-) 4-5 mm, lobes 
ovate to elliptic, obtuse, 4-7 mm, densely 
fimbriate; petals linear-lanceolate, 5-7 (-9) 
mm, 1-3-nerved, entire to densely fimbriate, 
inserted (1-) 2-8 mm above the corona; fila- 
ments 3-7 mm, connate up to 2,5 mm, inserted 
at the base of the hypanthium or on a short 
androgynophore; anthers 4—7 mm, obtuse; septa 
1-2,5 mm high; corona of fine hairs c. 1-2 mm; 
disc-glands 1-1,5 mm. Female flowers 
campanulate-infundibuliform, including the 
0, 5-2,5 mm long stipe (8-) 10-18 mm; hypan- 
thium tapering, 1-2 mm; calyx-tube 3-6 mm, 
lobes ovate, 4--7 mm, crenulate-laciniate; petals 
linear, acute, 3-5 (-6) mm, 1-nerved, entire or 
± fimbriate, inserted 1,5-3 mm above the 
corona; staminodes 2-2,5 mm; septa up to 0,2 
mm high; corona hairs 1-1,5 mm; disc-glands 
c. 0,5 mm; gynophore 1-1,5 mm; ovary ellip- 
soid to subglobose, (2-) 2, 5-4,5 mm; styles 
connate for 0,3-1, 5 mm, free style-arms 1,5— 
2,5 mm; stigmas palmately branched, papillate, 
each 2^2,5 mm diam. Fruit 1-2 per inflores- 
cence, subglobose, excluding the 1-3 mm long 
gynophore 2-3,5 (-6) x 2-3,5 (-5) cm; 
pericarp coriaceous, sometimes spongy inside; 
seeds 2-25 per capsule, ellipsoid, 6-7 (-8) mm. 

A species of savanna, stony slopes, rocky hillsides, 
stream banks, and forest along water-courses, found in 
Natal, Transvaal and Swaziland; also in southern Mozam- 
bique; 0-1200 m. Flowers in Aug.-Jan., fruits Nov.-Jan. 
Leaves without glands at the apex; glands at blade-base 
on a ± semi-orbicular appendage; petals in S 
flowers entire to remotely serrate-fimbriate, or 

fimbriate only at the base (a) var. hastata 

Leaves with glands at the apex; glands at blade-base 
on 2 separate auricles or on a bi-lobed append- 
age; petals in 6 flowers (mostly densely) 
fimbriate (b) var. glandulifera 


1 13 

Fig. 36. — 1, Adenia digitata, flowering stem, x % (Fries , N orlindh & Weimark 3415); a, female flower, longitudinal 
section, x 2Vi ( Chase 7887); b, male flower, longitudinal section, X 2'h (Liebenberg 3366); c, fruits, X ( Marques 

3443); d, seed, X 2Vi ( Munro s.n.); e-g, variation in leaf shape, all x V 2 : e (Young 33291), f ( Galpin 677), g (Munro 
s.n.) (after De Wilde, 1971). 


1 14 

(a) var. hastata. 

De Wilde in Meded. Landbouwhogeschool 
Wageningen 71-18: 170 (1971). 

Leaves without glands at the apex. 
Gland(s) at blade-base single, or 2 glands con- 
tiguous on a single, semi-orbicular, not lobed 
appendage. Male flowers 13-25 mm; petals 
entire or margin irregularly serrate-fimbriate, or 
fimbriate only at base; anthers in anthesis reach- 
ing to or nearly to the throat of the calyx-tube. 

The typical variety occurs in Natal, the eastern Trans- 
vaal lowveld, and in southern Mozambique, at 0-700 m 

Transvaal. — 2331 (Phalaborwa): Shingwedzi area. 
Van der Schijff 3850. 2431 (Acomhoek): Skukuza Camp, 
Codd 5732. 2531 (Komatipoort): Sigaas River, Van der 
Schijff 1305; Komatipoort, Schlechter 11747. 

Natal. — 2632 (Bela Vista): Ndumu Game Reserve, 
Ward 4520. 2732 (Ubombo): Pongola Poort, Strey 4643. 
2830 (Dundee): near Tugela Ferry, Dyer 4389. 2831 
(Nkandla): Mahlabatini, Gerstner 4234. 

Strey 4643, and probably also Codd 10281, both from 
northern Natal, possibly represent a separate form with 
large fruits up to 6 cm in diameter. These specimens have 
relatively large leaves, and were collected in forest along 

(b) var. glandulifera De Wilde in Meded. 
Landbouwhogeschool Wageningen 71-18: 170 
(1971). Type: Transvaal, Thorncroft 2034 

Leaves with 2 (-4)-paired, sometimes 
bulging glands at, or just below the apex. 
Glands at blade-base 2, on 2 separate auricles, 
or glands contiguous on the bi-lobed ± peltate 
blade-base. Male flowers 15-30 mm; petals 
moderately to densely fimbriate; anthers in 
anthesis remaining well below the throat of the 

Found in Natal, Swaziland and Transvaal, at 500-1200 
m altitude; it has a more western distribution than var. 

Transvaal. — 2530 (Lydenburg): near Nelspruit, Hunt 
Davy 1489; Leach 11540. 2531 (Komatipoort): Barberton, 
Galpin 563. 

Swaziland. — 2631 (Mbabane): Bulunga Poort, Comp- 
ton 31759. 

Natal. — 2731 (Louwsburg): near Nongoma, Acocks 
13019; Gerstner 2345; Pongola, Edwards 3188; Nel 215. 
2732 (Ubombo): Mkuzi, Venter 5204. 

The specimens Gerstner 2895, Edwards 3188, Nel 
215, and Compton 31759 from Natal and Swaziland are 
more or less intermediate between the 2 varieties, but in all 
4 the glands at the apex of the leaves are present. 

7. Adenia digitata (Harv.) Engl, in Bot. 
Jahrb. 14: 375 (1891); Burtt Davy in Ann. 

Transv. Mus. 3: 121 (1912); Engl., Pflanzenw. 
Afr. 3, 2: 605 (1921); Burtt Davy, FI. Transv. 1: 

221 (1926); Steyn, Tox. PI. S. Afr. 310, fig. 
43A, 44, 45, 47A (1934); Henkel, Woody PI. 
Natal 118 (1934); Liebenberg in Bothalia 3: 
541, 527, 530-532, fig. 1-3, 14-17, pi. 6-36 
(1939); A. & R. Fernandes in Garcia de Orta 6: 
259 (1958); Letty, Wild Flow. Transv. 225 
(1962); Watt & Breyer-Brandwijk, Med. Pois. 
PI. ed. 2: 826, fig. 218 (1962); De Wilde in 
Meded. Landbouwhogeschool Wageningen 
71-18: 188, fig. 30, 31 a-h (1971). Type: 
Zululand*, Owen s.n. (T.C.D., holo.; K, 
photo. !). 

Modecca digitata Harv., Thes. Cap. 1: 8, tab. 12 (1859); 
in F.C. 2: 500 (1862). M. senensis (Klotzsch) Mast, in 
F.T.A. 2: 513 (1871); Hook.f. in Bot. Mag. t.7763 (1901). 

Clemanthus senensis Klotzsch in Peters, Reise Mossamb. 
Bot. 143 (1862). Type: Mozambique, Peters s.n. (Bf, holo.; 
PRE, photo.!).. 

Adenia senensis (Klotzsch) Engl, in Bot. Jahrb. 14: 375 
(1891); Harms in Pflanzenfam. ed. 2, 21: 491 (1925); Bak. 
f. in J. Linn. Soc. (Bot.) 40: 73 (1910); Engl., Pflanzenw. 
Afr. 3, 2: 605 (1921); Burtt Davy, FI. Transv. 1: 221 (1926); 
A. & R. Fernandes in Garcia de Orta 6: 258 (1958). A. 
stenophylla Harms in Bot. Jahrb. 26: 238 (1899); Engl., 
Pflanzenw. Afr. 3, 2: 605 (1921); Burtt Davy, FI. Transv. 1: 

222 (1926). Type: Transvaal, Wilms 941 (Bt, holo.; PRE, 
photo.!). A. multiflora Pott in Ann. Transv. Mus. 5: 235 
(1917); Harms in Notizbl. Bot. Gart. Berl. 8: 298 (1923); 
Burtt Davy, FI. Transv. 1: 221 (1926). Type: Transvaal, 
Fehrson sub TRV 13786 (PRE, holo.!). A. angustisecta 
Burtt Davy in Kew Bull. 1921: 280 (1921); FI. Transv. 1: 
222 (1926). Type: Transvaal. Mundy 4700 (BOL, holo.; 
PRE, photo.!). A. buchananii Harms in Engl., Pflanzenw. 
Afr. 3, 2: 605 (1921). Lectotype: Malawi, Buchanan 244 
(K, lecto.) (see De Wilde, l.c.). 

(Sub)herbaceous climber 0,2-3 m growing 
from a variable tuber. Leaves beneath greyish 
to glaucous, densely punctate or not, deeply 
(3-) 5-parted or (pseudo-) foliolate, in outline 
suborbicular with cordate base, 4—18 x 3-17 
cm; leaflets variable, entire to deeply (2-) 3-5 
(-lO)-lobed, ovate or obovate to linear, base 
long-attenuate or acute, apex rounded to acute, 
1,5-15 X 0,7-4 (-7) cm, margin entire, 
petiolule up to 2 cm; petiole 1-9 cm. Glands at 
blade-base 2, on two separate ± upward di- 
rected auricles 1-2 mm diam. at the transition 
of petiole to blade; blade-glands 2-4, situated 
between the insertions of the leaflets, and each 
leaflet with 0-8 glands, scattered or submargi- 
nal. Stipules narrowly triangular to lanceolate, 
acute, 1-3 mm, withering. Inflorescences 

*It must be remembered that Miss Owen also collected 
near Rustenburg in the Transvaal and these gatherings were 
also placed as “Zululand” by Harvey. 



peduncled for up to 7 cm, (1-) 5-20 (-60)- 
flowered in 6, 1-10-flowered in 9 ; tendril 1, 

2- 10 cm, sterile tendrils up to 15 cm. Male 
flowers tubular-infundibuliform, including the 

3- 12 (-15) mm long stipe ( 14—) 20-38 mm; 
hypanthium cup-shaped, (1-) 2-3,5 x 2-4 (-5) 
mm; calyx-tube (5-) 8-12 mm; calyx-lobes 
ovate or oblong to lanceolate, (sub)obtuse, (4-) 
7-11 mm, dentate-fimbriate; petals lanceolate, 
acute, dentate or fimbriate, 6-12 mm, (1-) 
3-nerved, inserted at the same level as, or up to 
5 mm above the corona; filaments 3,5-9 (-12) 
mm, connate for at least half-way, inserted on 
an androgynophore 1-3,5 mm; anthers 3-6 mm, 
± inward curved and at the top clinging by the 
c. 0,2 mm long papillate apiculae; septa 0,5-3 
mm high; corona hairs c. 0,5-2 mm, rarely 
absent; disc-glands 0,5- 1,5 mm. Hermaphro- 
dite flowers 20-25 mm. Female flowers 
tubular-infundibuliform, including the 2-7 mm 
long stipe 15-26 mm; hypanthium c. 2-4 x 2-4 
mm; calyx-tube 4—8 mm, calyx-lobes ovate to 
oblong, obtusish, 5-7 mm, entire; petals lan- 
ceolate to linear, acute, (sub)entire, 2-7 mm, 1 
(-3)-nerved, inserted at the same level as, or up 
to 4 mm above the corona; staminodes 3-5 mm, 
up to 1 mm connate; septa up to 0,5 mm high; 
corona hairs 0,3-1 mm; disc-glands c. 1 mm; 
gynophore 2-4 mm; ovary ovoid to oblong, 
(4-) 5-6 mm; styles connate for 1-1,5 mm, 
style-arms 1—1,5 mm; stigmas subreniform, 
woolly-papillate, each 2-3 mm diam. Fruit 1-3 
per inflorescence, ovoid to ellipsoid (to oblong), 
excluding the (2-) 5-12 mm long gynophore 
(2,5-) 3-5,5 (-7,5) x (1,5-) 2-3,5 (-4) cm; 
pericarp coriaceous, brilliant yellow to red 
when fresh, sometimes spongy inside, smooth; 
seeds ( 10—) 20-60 per capsule, (ob)ovate to 
ellipsoid, 6-8 mm. Fig. 36. 

Found in Natal, Swaziland and Transvaal, outside the 
Flora area widely distributed in Angola, Tanzania, Zambia, 
Rhodesia, Malawi, Mozambique and Botswana. Grows in 
savanna, rocky and grassy places, on termite mounds, forest 
fringes, etc., in various soil types; 0-1850 m. Never 
gregarious. Flowers and fruits mainly Oct.-March. 

Transvaal. — 2229 (Waterpoort): near Ingwe Motel, 
Jacobsen 1905. 2230 (Messina): near Lake Funduzi, Story 
4856. 2231 (Pafuri): Baiandbai, Lang sub TRV 32154. 2327 
(Ellisras): Junction of Limpopo and Palala Rivers, Mogg 
24566. 2329 (Pietersburg): Breyer sub TRV 24215. 2330 
(Tzaneen): Westfalia Estate, Scheepers 515. 2428 

(Nylstroom): Mosdene, Galpin 13196. 2429 (Zebediela): 
Potgietersrus, Leendertz sub TRV 6007. 2430 (Pilgrims 
Rest): The Downs, Rogers sub TRV 18889. 2526 (Zeerust): 
Groot Marico, Liebenberg S.2. 2527 (Rustenburg): Brits, 
Mogg 14609. 2528 (Pretoria): near Rust de Winter, Codd 
2236. 2529 (Witbank): Loskop Dam, Theron 1943. 2530 

(Lydenburg): near Nelspruit, Schlieben 8449. 2531 

(Komatipoort): Barberton, Galpin 677. 2627 (Potchef- 
stroom): Experimental Farm, Theron 5. 2630 (Carolina): 
Lothair, Forester 28. 

Swaziland.— 263 1 (Mbabane): Hlatikulu, Ben Dlamini 

Natal. — 2632 (Bela Vista): Ndumu Game Reserve, 
Moll 4249.2130 (Vryheid): near Vryheid, Gerstner 2330. 
2731 (Louwsburg): Nongoma, Gerstner 4707. 2732 

(Ubombo): near Ingwavuma, Gerstner s.n. 2831 (Nkandla): 
6 km N. of Nkwaleni, A cocks 12954. 

This is an extremely variable taxon, but Liebenberg l.c. 
has convincingly shown that all forms belong to one single 

The fruits are poisonous to humans. 

8. Adenia wilmsii Harms in Bot. Jahrb. 
26: 238 (1899); Engl., Pflanzenw. Afr. 3, 2; 603 
(1921); Harms in Pflanzenfam. ed. 2, 21: 491 
(1925); Burtt Davy, FI. Transv. 1: 222 (1926); 
Liebenberg in Bothalia 3: 538, 526, 532, fig. 13 
(1939); Letty, Wild Flow. Transv. 225 (1962); 
De Wilde in Meded. Landbouwhogeschool 
Wageningen 71-18: 198 (1971). Type: Trans- 
vaal, Wilms 961 (Bt, holo.; BM; K; L; P; Z). 

Erect herb 10-30 (-50) cm, without ten- 
drils, glabrous, growing from a tuberous 
rootstock. Leaves ± glaucous-green, not 
punctate beneath, (5-) 7-pseudo-foliolate, 
broadly ovate to suborbicular in outline, base ± 
cordate, 2-8 (-12) x 2-6 (-10) cm, 5-7- 
plinerved; leaflets sessile, entire to pinnately 
3-7-lobed, ovate-oblong, base attenuate, apex 
acute, 1-6 (-10) cm, the middle leaflet largest; 
margin entire; petiole 2-8 (-10) cm. Glands at 
blade-base 2, on two auricles at the transition to 
the petiole; blade-glands (0— ) 4—6, situated at 
the very base in between the leaflets, sometimes 
a few submarginal. Stipules lanceolate, 1-3 
mm. Inflorescences peduncled for 0,5-5 cm, 
1-3-flowered in S, 1 (-3)-flowered in 9 ; no 
tendrils. Male flowers tubiform-infundibuli- 
form, including the 4—6 mm long stipe 20-25 
mm; hypanthium cup-shaped, 2-3 mm; calyx- 
tube 7-8 X 2-5 mm, lobes elliptic to oblong, 
obtuse, 6-7,5 mm, entire; petals oblanceolate, 
obtusish, 7-10 mm, 3-5-nerved, subentire, in- 
serted at the same level as, or up to 3 mm above 
the corona; filaments 4—5,5 mm, connate for 
1,5-2, 5 mm, inserted on a short an- 
drogynophore; anthers 3, 5^4, 5 mm; septa 2-3 
mm high; corona hairs 0,5-1 mm; disc-glands 
c. 0,5 mm. Female flowers tubiform- 
campanulate, including the 2-3 mm long stipe 
c. 12 mm; hypanthium c. 1 mm; calyx-tube 5-6 
X 3^4 mm, lobes elliptic-oblong, obtuse, c. 4 
mm, entire; petals oblanceolate, subobtuse, c. 6 



mm, 3-nerved, subentire, inserted at the same 
level as the corona; staminodes c. 2,5 mm; septa 
c. 1 mm high; corona hairs c. 0,5 mm; disc- 
glands c. 0,5 mm; gynophore 1,5 (-2) mm; 
ovary ovoid-ellipsoid, 4-4,5 mm; styles con- 
nate for 0,7 mm, style-arms c. 1 mm; stigmas 
subglobose, papillate, each c. 1,5 mm diam. 
Fruit 1 per inflorescence, ellipsoid, excluding 
the c. 5 mm long gynophore c. 4 x 2,5 cm; 
pericarp thinly coriaceous; seeds not known. 

This species is a local endemic in Transvaal, not found 
elsewhere, growing in rocky places, dolomite outcrops, or 
on red loam soil; at c. 1500 m. Flowers and fruits Sept.- 

Transvaal. — 2530 (Lydenburg): near Lydenburg, 

Liebenberg 3488; Strey & Schlieben 3116; Wilms 961; 
Van Wyk 7. 

9. Adenia repanda (Burch.) Engl, in Bot. 
Jahrb. 14: 375 (1891); Harms in Pflanzenfam. 
ed. 2, 21: 490, fig. 224 (1925); Liebenberg in 
Bothalia 3: 534, 525, 532, fig. 12 (1939); A. & 

R. Fernandes in Garcia de Orta 6: 658 (1958); 
C.F.A. 4: 219 (1970); Watt & Breyer- 
Brandwijk, Med. Pois. PI. ed. 2: 828 (1962); 
Schreiber in F.S.W.A. 89 (1968); De Wilde in 
Meded. Landbouwhogeschool Wageningen 
71-18: 237, fig. 37 (1971). Type: Cape, Griqua- 
land West, Burchell 2486/2 (K, holo.!; PRE). 

Paschanthus repandus Burch., Trav. 1: 543 (1822); 
Schinz in Bot. Jahrb. 15, Beibl. 33, 1:3 (1892); Marloth FI. 

S. Afr. 2, 2: 197, fig. 130 (1925). P. jdggii Schinz in Mem. 
Herb. Boiss. 20: 23 (1900). Type: as for Jaggia repanda 

Modecca paschanthus Harv. in F.C. 2: 500 (1862), nom. 
illegit. M. repanda (Burch.) Druce in Rep. Bot. Exch. Cl. 
Brit. Is. 636 (1917). 

Jaggia repanda Schinz in Verh. Bot. Ver. Prov. Bran- 
denb. 30: 254 (1888); Harms in Bot. Jahrb. 24: 169 (1897). 
Type: S.W. Africa, Schinz s.n. (Z, holo.!). 

Suberect herb or ± woody climber 0,2-2 
m, glabrous, growing from a tuberous 
rootstock. Leaves grey-glaucous, sometimes 
punctate beneath, entire or irregularly repand to 
lobed, obovate to linear, base attenuate to sub- 
cordate, apex subacute to obtuse or retuse, 
sometimes curved, 2-15 x 0,2-2 (-6) cm, ± 
penninerved; lobes up to 5 at either side of 
blade, up to 1 cm; petiole 0,1-1 cm. Glands at 
blade-base 2, one at each side of the base of the 
midrib; blade-glands 0-10, submarginal, mostly 
corresponding with the lobes, and one apical or 
subapical gland in which the midrib ends. 

Stipules narrowly triangular, acute, 1-1,5 mm. 
Inflorescences sessile or peduncled for up to 2 
cm, 1-5-flowered in S, 1-2 (-3)-flowered in 5 
and $; tendril (0 or) 1, up to 4 cm; sterile 
tendrils up to 5 cm; flowers polygamous or 
dioecious. Male flowers tubiform- 
infundibuliform, including the 2-3 (-4) mm 
long stipe 15-24 mm; hypanthium including 
calyx-tube 9-14 x 2-5 mm; calyx-lobes 
oblong-lanceolate, obtuse, 4—7,5 mm, entire; 
petals lanceolate, obtuse to subacute, 5-8 mm, 
(1-) 3-nerved, subentire, inserted near the 
throat of the calyx- tube, 8-12 mm above the 
base of the hypanthium; filaments 3,5-6 mm, 
free, inserted in the calyx-tube 4—5 mm above 
the base of the hypanthium; anthers 4—6 mm; 
septa 0; corona 0; disc-glands 0. Hermaphro- 
dite flowers ± intermediate between male and 
female flowers; anthers smaller. F emale flowers 
tubiform-campanulate, including the c. 1 mm 
long stipe 8-11 mm; hypanthium including 
calyx- tube 3,5-5 x 3-4 mm; calyx-lobes ob- 
long to lanceolate, obtuse, 3-5 mm, entire; 
petals lanceolate, obtuse or acute, 1,5-2 mm, 1 
(-3)-nerved, entire, inserted 2, 5-3, 5 mm above 
the base of the hypanthium; staminodes 2-3 
mm, inserted near the base of the hypanthium; 
septa 0; corona 0; disc-glands 0; gynophore 2-3 
mm; ovary ovoid to ellipsoid, 2,5-4 mm; styles 
connate for 0,5-1, 5 mm, style-arms 0,5-0, 7 
mm; stigmas subglobose, papillate, each 1-1,5 
mm diam. Fruit 1-2 per inflorescence, sub- 
globose, excluding the 2-5 mm long gynophore 
1,5-2, 5 (-3) x 1, 2-2,5 cm; pericarp coriace- 
ous, sometimes spongy inside; seeds (2-) 5-12 
per capsule, suborbicular to broadly ovate, 7-8 

This species, which represents a monotypic section 
Paschanthus in Adenia, is distributed in rather restricted 
areas in Southern Africa, viz. the northern Cape Province, 
the Soutpansberg area in the Transvaal, and throughout 
South West Africa; it also occurs in Angola, Zambia, 
Rhodesia, and Botswana. The species is often recorded as 
“rare” or “uncommon”, and grows in open or partly 
shaded places in woodland, among rocks, on sandy soils 
(“Kalahari sand”), red sand, and granite; 500-1 600 m. 
Flowering time Oct.-Febr., fruits Nov.-April. 

S.W.A. — 1715 (Ondangua): Ondangua, De Winter & 
Giess 6853. 1817 (Tsintsabis): near Tsintsabis, Boss sub 
TRY 35549. 1917 (Tsumeb): Kududamm, Volk 413. 1920 
(Tsumkwe): Gautscha Pan, Story 6284. 2117 (Otjosondu): 
Quickbom, Bradfield 75. 2217 (Windhoek): Windhoek, 
Rogers 29798. 2718 (Griinau): Karasberg, Pearson 8166. 

Transvaal. — 2229 (Waterpoort): between Soutpan and 
Waterpoort, Obermeyer, Schweickerdt & Verdoorn 246. 
2230 (Messina): Tshipise, Van der Schijff 5215. 2329 
(Pietersburg): Vivo, Bremekamp & Schweickerdt 206. 


1 17 

Cape. — 2624 (Vryburg): Armoedsvlakte, Mogg 8769. 
2723 (Kuruman): near Kuruman, Marloth 1350. 2824 
(Kimberley): Barkly West, Acocks 1561. 2922 (Prieska): 
near Prieska, Bryant 345. 

According to Bryant 345 “greedily eaten by stock, 
which may partly account for its rarity”, but also reported 
as “deadly poisonous to man”. 

10. Adenia gummifera (Harv.) Harms in 
Pflanzenfam. 3, 6a, Nachtr. 1: 255 (1897); ed. 
2, 21: 490 (1925); Burtt Davy in Ann. Transv. 
Mus. 3: 121 (1912); FI. Transv. 1: 222 (1926); 
Henkel, Woody PI. Natal 110 (1934); Lieben- 
berg in Bothalia 3: 535, 523, 532, fig. 10 & 1 1 
(1939); A. & R. Fernandes in Garcia de Orta 6: 
256 (1958); Watt & Breyer-Brandwijk, Med. 
Pois. PI. ed. 2: 828 (1962); F. White, For. FI. N. 
Rhod. 267 (1962); Letty, Wild Flow. Transv. 
225 (1962); De Wilde in Acta Bot. Neerl. 17: 
131, fig. 2h (1968); in Meded. Land- 
bouwhogeschool Wageningen 71-18: 261 

(1971). Type: South Africa, “inter Omsamculo 
et Omcomas”, Drege 5211 (P, holo.!; S). 

Modecca gummifera Harv. in F.C. 2: 500 (1862). 

Ophiocaulon gummifer (Harv.) Mast, in F.T.A. 2: 518 
(1871); Harms in Engl., Pflanzenw. Ost Afr. 2, C: 281 
(1895) ( as gummiferum ); Fries, Wiss. Ergebn. Rhod. Kongo 
157 (1914). O. cissampeloides sensu Bak.f., FI. Maur. and 
Seychelles 106 (1877); in J. Linn. Soc. (Bot.) 40: 74 ( 1910). 

Adenia rhodesica Suesseng. in Trans. Rhod. Sc. Ass. 43: 
13 (1951). Type: Rhodesia, Dehn 696/52 (M, holo.!; BM; 

Subligneous climber up to 30 m, up to 10 
cm thick at base; twigs green or grey-green, 
often pruinose. Leaves rarely punctate, entire to 
deeply 3 (-5)-lobed, orbicular to ovate or 
rhomboid, or ± 3 (-5)-angular in outline, base 
cordate to truncate, apex obtuse or retuse, rarely 
subacute, 2,5-1 1 x 2,5-1 1 cm, 3-plinerved and 
with 1 pair of straight nerves from the midrib 
ending in marginal glands; reticulation rather 
fine, distinct; margin entire; lobes obtuse, up to 
4 cm; petiole 1,5-11 cm. Gland at blade-base 
single, on a median subcircular to spathulate 
appendage 1-3 mm; blade-glands 0-4, often 2 
glands rather approximate to the axils of or 
contiguous with the upper side nerves; marginal 
glands minute, 3-7 on either side of the blade. 
Stipules broadly rounded to triangular, finely 
lacerate, 0,5 (-1) mm. Inflorescences pedun- 
cled for (0,5—) 1-12 (-16) cm, up to 35- 
flowered in d>, 2-6-flowered in ?; tendrils 0 or 1, 

1- 4 cm, sterile tendrils simple or 3-fid, 5-20 
cm. Male flowers ± campanulate, including the 

2- 6 mm long stipe 11-17 mm; hypanthium 

cup-shaped, 1-2 (-2,5) x 2-4 mm; calyx-tube 
0; sepals lanceolate, subobtuse, (7-) 8-10 x 
2-3 mm, margin up to 0,2 mm laciniate, 
punctate; petals (ob)lanceolate, obtuse, (6-) 
8-11 mm, 3-nerved, finely laciniate-serrulate, 
remotely punctate; filaments (1-) 2-3,5 mm, 
connate for (0-) 0,5-1, 5 (-2) mm, inserted at 
the base of the hypanthium; anthers 3-6 mm, 
obtuse, up to 0, 1 mm apiculate; septa 0-0,2 mm 
high; corona 0; disc-glands 0. Female flowers 
± campanulate, including the c. 0,5 mm long 
stipe 5,5-8 mm; hypanthium flatfish, c. 0,5 x 
2-2,5 (-3) mm; calyx-tube 0; sepals oblong, c. 
4—6,5 mm, entire, punctate; petals lanceolate- 
linear, 2-4,5 mm, 1-3-nerved, sub-entire, spar- 
ingly punctate or not; staminodes c. 0,5 mm; 
septa 0; corona 0; disc-glands 0; gynophore c. 
0,5 mm; ovary ovoid, 3-4,5 mm; style 0-0,5 
mm; stigmas (sub)sessile, subreniform, 
laciniate-papillate, each c. 1-1,5 mm diam. 
Fruit 1-4 per inflorescence, ovoid to ellipsoid, 
sometimes ± 3 (-6)-angular, excluding the c. 1 
mm long gynophore 2,5-4 (-4,5) x 1,7-3 cm; 
pericarp woody-coriaceous, c. 0,2 mm thick, 
smooth or finely pitted or granulate; seeds 
30-50 per capsule, subovate, 3, 5-5, 5 mm. 

Widely distributed in eastern Africa, from Somalia in 
the north to the Kei River in the south; also in Seychelles. It 
is a locally common, vigorous trailer, found in forest and 
scrub at 0-1 800 m. Flowering time mainly Sept.-April. 

Transvaal. — 2229 (Waterpoort): 30 km N. of Louis 
Trichardt, Buitendag 1075. 2230 (Messina): Pepiti Falls, 
Smuts & Gillen 3221. 2231 (Pafuri): Punda Milia, Codd 
5966. 2329 (Pietersburg): 8 km W. of Louis Trichardt, 
Obermeyer, Schweickerdt & Verdoorn 355. 2330 (Tza- 
neen): Westfalia Estate, Scheepers 744. 2429 (Zebediela): 
Sekukuniland, Barnard 128. 2430 (Pilgrims Rest): 

Mariepskop, Van der Schijff 4736. 2431 (Acornhoek): 
Lothian, Strey 3573. 2530 (Lydenburg): Sabie, Louw 2398. 
2531 (Komatipoort): Barberton, Galpin 782. 

Swaziland. — 2631 (Mbabane): near Mbabane, Com- 
pton 30370. 

Natal. — 2632 (Bela Vista): Kosi Bay, Strey & Moll 
3829. 2732 (Ubombo): Sordwana Bay, Vahrmeijer & 
Tolken 314. 2830 (Dundee): near Tugela Ferry, Dyer 4389. 
2831 (Nkandla): Eshowe, Gerstner 2641. 2832 

(Mtubatuba): Hluhluwe Game Reserve, Ward 3962. 2930 
(Pietermaritzburg): Berea, Durban, Medley Wood 5502. 
2931 (Stanger): Hawaan Forest, Ross & Moll 2265. 3030 
(Port Shepstone): Dumisa, Rudatis 523. 

Cape. — 3129 (Port St. Johns): Port St. Johns, Galpin 
3461 . 3228 (Butterworth): Kentani, Pegler 869. 

In South Africa the typical variety, var. gummifera 
occurs; specimens from a restricted area in Zambia have 
been described as var. cerifera (De Wilde, l.c. p. 264). 




Schlechterina Harms in Bot. Jahrb. 33: 148 (1902); in Ber. Deutsch. Bot. Ges. 24: 177, 1. 12 
(1906); in Engl., Pflanzenw. Afr. 3, 2: 596 (1921); Pflanzenfam. ed. 2, 21: 485, fig. 221 (1925); 
Marloth, FI. S. Afr. 2, 2: 198 (1925); Ross, FI. Natal 251 (1972); De Wilde in Blumea 22: 48 
(1974). Type: S. mitostemmatoides Harms. 

Crossostemma sensu Hutch., Gen. FI. Pi. 2: 370 (1969), pro parte. 

Low perennial climber or suberect shrublet, mostly provided with tendrils, glabrous, growing 
from a perennial rootstock. Leaves simple, elliptic to linear, entire to deeply pinnately lobed, 
shortly petiolate, margin (sub)entire to dentate. Glands on petiole 0 or 1 (or 2) pairs at the top. 
Stipules small, caducous. Tendrils axillary. Inflorescences axillary, (sub)sessile, few-flowered, 
often arranged in short shoots from the supra-axillary bud; bracts and bracteoles small. Flowers 
hermaphrodite, whitish; stipe articulate to short pedicel; hypanthium small, shallowly cup-shaped. 
Sepals 3 or 4, free, imbricate, elliptic to oblong. Petals 2-4, free, elliptic to oblong. Corona single, 
composed of threads connate at base into a low tube, inside set with additional hair-like 
appendages. Disc absent. Androgynophore short. Stamens 6-8, connate at base into a shallow cup, 
often with small lobes (staminodes) on its margin in between the filaments; anthers dorsifixed, 
versatile, ellipsoid-oblong, obtuse, 2-locular. Ovary ellipsoid-oblong, on a short gynophore; 
placentas (3 or) 4, each with 3-8 ovules; style single; stigma single, flatfish, 3- or 4-lobed. Fruit a 
stipitate 3- or 4-valved capsule, ellipsoid-oblong, fusiform; valves coriaceous. Seeds flattened, 
ellipsoid, arillate; testa crustaceous, scrobiculate. 

One species in tropical East Africa, just entering northern Natal. 

Related to the West African genus Crossostemma, but the latter is distinguished by a conspicuous intra-staminal disc, 
and the absence of a gynophore. 

Schlechterina mitostemmatoides Harms 
in Bot. Jahrb. 33: 148 (1902); in Ber. Deutsch. 
Bot. Ges. 24: 111 (text fig.), 1. 12 (1906); Bak. f. 
in J. Linn. Soc. (Bot.) 40: 73 (1911); Engl., 
Pflanzenw. Afr. 3, 2: 596 (1921); Harms in 
Pflanzenfam. ed. 2, 21: 485, fig. 221 (1925); A. 
& R. Fernandes in Garcia de Orta 6: 249 
(1958); Ross, FI. Natal 251 (1972). Type: 
Mozambique, Lourenco Marques, Schlechter 
1 1681 (B, holo.f). 

S. mitostemmatoides var. holzii Harms in Ber. Deutsch. 
Bot. Ges. 24: 184 (1906); Engl., Pflanzenw. Afr. 3, 2: 596 
(1921); Harms in Pflanzenfam. ed. 2, 21: 486 (1925). 
Syntypes: Tanzania, near Dar-es-Salaam, Holz 1070, 
1070a; 1086 (Bf). 

Small liana or subscandent shrublet to 3 m, 
glabrous, with perennial rootstock; older stem 
corky, shoots often lenticellate; tendrils 3-10 
(-15) cm. Leaves simple, variable 
(heterophyllous), elliptic to lanceolate (to 
linear), apex acute to (long) acuminate, base 
acute to attenuate, margin entire or regularly to 
irregularly dentate, or leaves pinnately lobed to 
various depth, the lobes acute to rounded, 
2,5-13 x 0,7-4, 5 cm; leaves in sapling-, or 

sterile- or juvenile shoots often lanceolate to 
linear, (deeply) pinnately lobed, the lobes broad 
or narrow, 10-30 x 0,2-2 cm; petioles 4—12 
mm, in juvenile leaf-forms 0-4 mm; glands on 
petiole absent, or 1 (or 2) pair(s) at the apex; 
glands on blade-margin several, minute, mostly 
at the tips of the teeth; stipules subtriangular- 
linear, c. 0,5 mm. Inflorescences 1-3-flowered, 
often arranged in short shoots from the supra- 
axillary bud; bracts subtriangular, 0,5-1 mm. 
Flowers glabrous, stipe 6-25 mm; hypanthium 
small, shallowly cup-shaped, c. 3-4,5 mm 
wide; sepals 3 or 4, elliptic to oblong, obtuse, 
6-11 x 3-6 mm; petals 2-4, elliptic to oblong, 
obtuse, 5-10 mm long; corona single 5-8 mm 
high, composed of threads ± connate at base 
into a tube 0,5-2 mm, free parts of threads 4—6 
mm, inside tube and at base of free threads a 
zone of short hair-like appendages 0,5-1 mm. 
Disc absent. Androgynophore c. 1 mm. Sta- 
mens 6-8 (often 7), 7-10 mm, connate at base 
into a cup c. 1 mm, often with small lobes on its 
margin in between the filaments; anthers 
ellipsoid-oblong, obtuse, 2-3 mm. Gynophore 
2-2,5 mm; ovary ellipsoid-oblong, 2-2,5 mm, 
± (3- or) 4-angled; style single, 1,5-2 mm; 



Fig. 37. — Schlechterina mitostemmatoides, flowering stem, x 2 h\ a, flower, x 3 (Faulkner 1773); b-e, variation in leaf 
shape, all X 2 h: b (Strey 10439), c (Pedro 328), d (Sim 20660), e (/Marques 2242). 



stigma single, 3- or 4-lobed, 2-2,5 mm diam. 
Fruit a (3- or) 4-valved capsule, ellipsoid- 
oblong, acute at both ends, excluding the c. 1 
cm long gynophore 4,5-5 x 2,5-3 cm; seeds 
rather few, ellipsoid, c. 8 mm. Fig. 37. 

This species enters the Flora area in the very north-east 
of Natal, growing in coastal scrub and forest. Its main area 

of distribution is Kenya, Tanzania, Zanzibar and Mozam- 
bique, where it is recorded from sandy soils and old coral 
reefs, at an altitude from 0-700 m. 

Natal. — 2632 (Bela Vista): 8 km N.E. of Makanes Pont, 
Ross 2370; Strev 10439. 2732 (Ubombo): Makanes Pont, 
Venter 5122. 

The species displays a remarkable variability in leaf 
shape (see description). 


Basananthe Peyr. in Bot. Ztg. 17: 101 (1859); (in Wawra & Peyr., Sertum Benguelense in) Sitz. 
Ber. Acad. Wien 38: 569 (1860); Hook.f. in Benth. & Hook.f., Gen. PI. 1: 812 (1867); Welw. in 
Trans. Linn. Soc. Lond. 27: 27 (1871); Mast, in F.T.A. 2: 508 (1871); Trans. Linn. Soc. Lond. 27: 
639 (1871); Hiem, Cat. Afr. PI. Welw. 1, 2: 382 (1898); De Wilde in Blumea 21, 2: 327 (1974). 
Type: B. littoralis Peyr. 

Tryphostemma Harv., Thes. Cap. 1: 32, t. 5 1 (1859); in F.C. 2: 499 (1862); Hook.f. in Benth. & Hook.f., Gen. PI. 1:811 
(1867); Mast, in F.T.A. 2: 507 (1871); Engl, in Bot. Jahrb. 14: 387 (1891) (including sect. Eutryphostemma, sect. 
Neotryphostemma, and sect. Basananthe)-, Pflanzenw. Afr. 3, 2: 598 (1921); Harms in Notizbl. Bot. Gart. Berl. 8: 291 
(1923) (including ‘Reihen 'Appendiculatae and Exappendiculatae); in Pflanzenfam. ed. 2, 21: 487 (1925); Hutch. & Pearce 
in Kew Bull. 1921: 257 (1921) (including series Lobatifoliae and Integrifoliae); Phill. , Gen. ed. 2: 518 (1951); A. & R. 
Fernandes in Garcia de Orta 6: 250, 662 (1958); Hutch., Gen. FI. PI. 2: 371 (1967); Ross, FI. Natal 251 (1972). 

Carania Chiov., FI. Somala 175 (1929). 

Annual or perennial herbs or small climbers, rarely shrubs, glabrous or hairy, with or without 
tendrils. Leaves simple (not lobed) or lobed, sessile or petiolate; margin entire or mostly dentate 
with small glandular teeth; apex mostly mucronate. Stipules small, linear; false stipules in some 
species, developed from the supra-axillary bud. Tendrils axillary, replacing central flower of cyme, 
or absent. Inflorescences axillary, cymose, 1-3-flowered, sessile or peduncled; bracts and 
bracteoles small, linear, often forming an involucre. Flowers bisexual (sometimes functionally 
unisexual), campanulate, greenish; stipe indistinctly articulate at base to the short pedicel; 
hypanthium rather narrow, flattish, rarely shallowly cup-shaped. Sepals 5 (or 6), oblong to 
lanceolate, free. Petals absent, or 1-2, or (4 or) 5 (or 6), oblong to lanceolate, (sub)obtuse, free, 
usually smaller than the sepals. Outer corona consisting of a ± barrel-shaped tube bearing a ring of 
filiform processes (threads), bluish, and mostly with a ring of small inward curved teeth. Disc low, 
annular, rarely absent. Inner corona membranous, cup-shaped, margin entire or lobulate, in B. 
berberoides from Somalia forming 5 small cups around the bases of the filaments. Stamens 5 (or 
6-9); filaments inserted in the upper half inside the corona, free; anthers basifixed, ellipsoid to 
lanceolate, (sub)sagittate, 2-locular. Ovary ellipsoid, superior, mostly sessile, 1-celled, with 3 (or 
4) placentas; styles 3 (or 4), free or partially united; stigmas globose, small. Fruit a sessile or 
shortly stipitate 3- (or 4)-valved capsule, ellipsoid; valves coriaceous. Seeds 1 or a few, arillate, 
ellipsoid to reniform, ± compressed; testa coriaceous, mostly rugose, blackish. 

A genus of 25 species in central, east and southern Africa; 5 species occur in the Flora area. 

As I have explained in my revision (l.c. ), I have refrained from subdividing the genus into sections. 

False stipules, as used in key and descriptions, are stipule-like appendages much larger than the minute true stipules, 
developed from the supra-axillary serial bud or (short) shoot. 

False stipules large, foliaceous; petals present: 

Leaves 3-9-lobed: 

Sepals 3-4,5 mm, with keeled nerves; leaves 3-7 (~9)-lobed 1. B. pedata 

Sepals 5-7,5 mm, not keeled; leaves 3-lobed 2. B. triloba 

Leaves simple, rarely partly faintly lobed 3. B. polygaloides 

False stipules absent, or the first leaves (cataphylls) of supra-axillary shoot not distinctly appearing as false stipules; 

petals present or absent: 

Petals absent, or 1 or 2; leaves not lobed 4. B. sandersonii 

Petals present, 4 or 5; leaves, at least in part, deeply 3-lobed 5. B. heterophylla 



Fig. 38. — 1, Basananthe triloba, flowering stem, x 1; la, flower, x 4 {Pott 5454); lb, leaf, X 1 ( Compton 31222); 2. B. 
sandersonii, leaf, false stipules absent, x 1 (Acocks 1 1795); 3, B. polygaloides, leaf and false stipules, x 1 (garrison 
296); 4, B. pedata, leaf and false stipules, x 1 (Wild & Drummond 7001) (after De Wilde, 1973). 



1. Basananthe pedata (Bak.f.) De Wilde 
in Blumea 21: 333, fig. la, 5 (1974). Type: 
Rhodesia, Rand 67 (BM). 

Tryphostemma pedarum Bak.f. in J. Bot. Lond. 37: 436 
(1899); Hutch. & Pearce in Kew Bull. 1921: 263 (1921). T. 
schlechteri Schinz in Vjschr. Naturf. Ges. Zurich 55: 243 
(1911); Schreiber in F.S.W.A. 89: 2 (1968). Type: Trans- 
vaal, Schlechter 4596 (B, holo. !; HBG). T. arenophilum 
Pott in Ann. Transv. Mus. 5: 234 (1915). Type: Transvaal, 
Leendertz 2062 (PRE, holo.!; K). T. harmsianum Dinter in 
Feddes Rep. 24: 304 (1928). Type: S.W. Africa, Dinter 872 
(Bf, holo.; M). 

Erect annual or biennial herb up to 50 cm, 
glabrous or slightly scabrous, often with some 
spreading-erect branches from the base; tendrils 
absent. Leaves deeply pedately 3-7 (-9)-lobed, 
the lobes elliptic to linear, (0,5-) 1-8 cm, 
obtuse to acute-acuminate, margin remote- to 
densely dentate-mucronate (serrate) up to 1,5 
mm; leaf-base subcordate to long-cuneate, de- 
current in the alate petiole 0,2- 1,5 cm. Stipules 
2-6 mm; false stipules foliaceous, asymmetri- 
cal, 0,5-2 cm, acute-acuminate, mucronate. 
Inflorescences 1- or 2-flowered; peduncle up to 
2 cm; bracts 2-4 mm. Flowers glabrous; stipe 

1.5- 3, 5 mm; hypanthium 1-2 mm wide. Sepals 

2. 5- 4,5 mm, with (2-) 3-keeled or winged 
green nerves. Petals 2-4,5 mm. Outer corona- 
tube (0,5-) 0,7-1, 3 mm, threads (0,5-) 0,7-1 
mm. Disc 0, 1-0,2 mm. Inner corona cup- 
shaped, 0,3-0, 8 mm. Stamens 5; filaments 
1-1,5 mm; anthers 0,5-1 mm. Ovary 0,5-1, 5 
mm; styles free, 0,6-2, 3 mm. Fruit subsessile, 
(0,7-) 1 cm, containing 1 seed c. 7 mm. Fig. 38: 

Growing mostly in sandy soil of riverbeds or dunes, or 
in gravelly, or sometimes rocky soil; 400-1 200 m; occurs 
locally in Transvaal and in north-eastern South West Africa; 
also in Zambia, Rhodesia and Botswana. 

S.W. A. — 1716 (Enana): near Oshandi, De Winter <& 
Giess 7027. 1718 (Kuring-kuru): Omuramba Mpungu, De 
Winter 3902. 1721 (Mbambi): Shamvura, De Winter & 
Marais 4600. 1819 (Karakuwisa): Karakuwisa, Dinter 
7269. 1820 (Tarikora): 16 km N. of Tamso, De Winter & 
Marais 4727. 

Transvaal. — 2329 (Pietersburg): Kalkbank, Acocks 
8848. 2428 (Nylstroom): near Warmbaths, Leendertz 2062. 
2528 (Pretoria): 14 km E. of Hammanskraal, Meeuse 9573. 

2. Basananthe triloba ( H . Bol.) De 
Wilde in Blumea 21: 335, fig. lb, 5 (1974). 
Type: Mozambique, Delagoa Bay, Bolus 7606 
(K, holo.!; BM). 

Tryphostemma trilobum H. Bol. in Hook. Icon. PI. 19: 
1. 1838 (1889); Schinz in Bot. Jahrb. 15, Beibl. 33: 3 (1892); 
Hutch. & Pearce in Kew Bull. 1921: 261 (1921); Harms in 
Pflanzenfam. ed. 2, 21: 488 (1925); A. & R. Fernandes in 

Garcia de Orta 6: 252 (1958). T. schinzianum Harms in 
Pflanzenfam. 3, 6a: 72, 75, fig. 25a, 26c (1893); ed. 2, 21: 
488, fig. 217c (1925); in Engl., Pflanzenw. Ost Afr., C: 281 
(1895); Bak.f. in J. Linn. Soc. (Bot.) 40: 73 (1911); Hutch. 
& Pearce in Kew Bull. 1921; 263 (1921); A. & R. 
Fernandes in Garcia de Orta 6: 253 (1958). Type: Mozam- 
bique, Stuhlmann 835 (Bf, holo.; BM, drawing; HBG). T. 
sagittatum Hutch. & Pearce in Kew Bull. 1921: 262 (1921); 
Harms in Pflanzenfam. ed. 2, 21: 488 (1925). Syntypes: 
Transvaal, Galpin 505 (PRE!); Kirk 74 (K!). 

Perennial herb, several shoots creeping or 
climbing to 70 (-250) cm, branched or not from 
a woody rootstock, glabrous; tendrils up to 10 
cm. Leaves ± sagittate, 3-lobed, 1,5-8 x 1-7,5 
cm, the lobes suborbicular to oblong, up to 5,5 
x 3 cm, apex obtuse to acute, the middle lobe 
sometimes largest; margin simple or double 
(densely) serrate-dentate 1-2 mm; petiole 0, 1-4 
cm. Stipules 3-8 mm; false stipules ± asym- 
metrical, 2-lobed, serrate-mucronate, 0,5-3, 5 x 
0,2-1 cm. Inflorescences (1- or) 2-flowered; 
peduncle 1-6 cm, bracts 2-3 mm. Flowers 
glabrous; stipe 4-7 mm; hypanthium 2,5-3 mm 
wide. Sepals 5-7,5 mm. Petals 4-6,5 mm. 
Outer corona-tube 1,7-2, 5 mm, threads 0,5- 1,3 
mm. Disc c. 0,2 mm. Inner corona cup-shaped, 
1,2-2 mm. Stamens 5; filaments 2-2,5 mm 
(sometimes alternating with small tooth-like 
appendages); anthers 1,2-1, 8 mm. Ovary up to 
1 mm stipitate, 1-1,5 mm; styles (1-) 2,5-4 
mm, free or up to 1 mm connate. Fruit (exclud- 
ing the 2-4 mm long gynophore) c. 1,2-1, 5 cm, 
containing 2 or 3 seeds c. 7 mm. Fig. 38: 1. 

Distributed in Natal, Swaziland and Transvaal, extend- 
ing to the north in Mozambique and Rhodesia; found on 
sandy or stony soil, in grassland, scrub, and open forest; 
0-1 200 m. Flowers and fruit mainly Sept.-Febr. 

Transvaal. — 2431 (Acomhoek): Klaserie, Killick & 
Strey 2515. 2530 (Lydenburg): Lowveld Botanic Garden, 
Buitendag 214. 2531 (Komatipoort): near Pretorius Kop, 
Codd 5668 ; Barberton, Galpin 505. 

Swaziland. — 2631 (Mbabane); near Stegi, Compton 
28394; 31222. 

Natal. — 2732 (Ubombo): near Jozini, Strey 10250. 
2831 (Nkandla): near Hlabisa, Gerstner 5022. 

3. Basananthe polygaloides (Hutch. & 
Pearce) De Wilde in Blumea 21: 338, fig. Id, 5 
(1974). Type: Natal, Medley Wood 10339 (K, 
holo. !). 

Tryphostemma polygaloides Hutch. & Pearce in Kew 
Bull. 1921: 263 (1921); Harms in Pflanzenfam. ed. 2, 21: 
488 (1925); Ross, FI. Natal 251 (1972). 

Perennial herb, up to 100 cm, with woody 
rootstock, glabrous; tendrils 2-8 (-10) cm, or 
none. Leaves simple or rarely faintly 3-lobed, 



ovate to elliptic, 2-7,5 x 1-5 cm, base subcor- 
date to acute, apex subacute to acute-acuminate, 
margin single- or double serrate-dentate 1-3 
mm deep; petiole 0, 1-0,5 cm. Stipules 2,5-8 
mm; false stipules asymmetrical, sharply den- 
tate, 0,4-1, 5 cm. Inflorescences (1- or) 

2- flowered; peduncle 1-5,5 cm, bracts 2-6 mm. 
Flowers glabrous; stipe 5-15 mm; hypanthium 

3- 4,5 mm wide. Sepals 7-10 mm. Petals 6,5-9 
mm. Outer corona-tube 2-2,5 mm, with a 
plicate, inward-folded edge; threads 1-2 mm. 
Disc c. 0,2 mm. Inner corona cup- or funnel- 
shaped, 1,5-2 mm. Stamens 5; filaments 2,5- 

4.5 mm (sometimes alternating with small 
tooth-like appendages); anthers c. 2 mm. Ovary 

1.5- 2mm; styles 3, 5-4, 5 mm, connate for 0,5- 

2.5 mm. Fruit (excluding the 2-4 mm long 
gynophore) 1,5—2, 2 cm, containing 2-4 seeds 
8-9 mm. Fig. 38: 3. 

A local endemic of South Africa, in Zululand (Natal), 
not occurring elsewhere. Grows in grassland, bushes, and 
coastal forest, always reported from sandy soil; 0-200 m. 
Flowers and fruits Jan. -March, and Sept.-Nov. 

Natal. — 2732 (Ubombo): Mpangazi, Strey 5055. 2832 
(Mtubatuba): 26 km N. of Mtubatuba, Acocks 13082; near 
Cape Vidal, Codd 10166 ; Richards Bay, Venter 4867. 

4. Basananthe sandersonii (Harv.) De 
Wilde in Blumea 21: 339, fig. 2a-c, 5 (1974). 
Type: Natal, Sanderson 1864 (TCD, holo.;.K). 

Tryphostemma sandersonii Harv., Thes. Cap. 1: 33, t.51 
(1859); in F.C. 2; 499(1861); Hutch. & Pearce in Kew Bull. 
1921: 265 (1921); Ross, FI. Natal 251 (1972). T. natalense 
Mast, in Trans. Linn. Soc. Lond. 27: 639 (1871), nom. 
illegit. Type as for T. sandersonii. T. longifolium Harms in 
Bot. Jahrb. 33: 149 (1902); Hutch. & Pearce in Kew Bull. 
1921: 264 (1921); A. & R. Fernandes in Garcia de Orta 6: 
252 (1958). Type: Tanzania, Basse 673 (Bf, holo.; EA). T. 
viride Hutch. & Pearce in Kew Bull. 1921: 265 (1921); 
Ross, FI. Natal 251 (1972). Syntypes: Transvaal, Bolus 
7602 (K!); Galpin 931 (PRE!; K!); Thorncroft 4366 (K!). 
T.friesii Norlindh in Bot. Notiser 107, fig. 9, 10 (1934); A. 
& R. Fernandes in Garcia de Orta 6: 251, 252 (1958). Type: 
Rhodesia, Fries, Norlindh & Weimarck 3112 (S, holo.!; 
BM!; BR!; PRE!). 

Perennial herb 2-60 cm, glabrous, 1 to 
several shoots erect or ± prostrate at base from 
a rootstock; tendrils sometimes present, 0,5-2 
(-5) cm. Leaves simple, 2-16 x 0,7-4 (-5,5) 
cm, suborbicular or broadly ovate (or obovate) 
or elliptic to lanceolate; base subcordate or 
rounded to acute (to attenuate), apex broadly 
obtuse (to retuse) to (mostly) acute, margin 
remotely serrate-dentate 0,5-1 mm, especially 
towards base; petiole 0-0,3 (-0,5) cm. Stipules 

1.5- 5 mm; false stipules absent. Inflorescences 
1-3-flowered; peduncle up to 4,5 cm; bracts 

1-4 mm. Flowers glabrous; stipe 3-17 mm; 
hypanthium 2-4 mm wide. Sepals 5-7, (the 
inner ones petaloid), obtuse, 4, 5-8, 5 mm. Pe- 
tals 0 (-2). Outer corona-tube 1-2 (-3) mm, 
threads 0,7- 1,5 (-2) mm, sometimes ± 
branched. Disc 0, 1-0,5 mm. Inner corona 
cup-shaped 1-1,5 (-2,5) mm. Stamens 5; fila- 
ments 1,5-3, 5 mm, anthers 0,7- 1,3 mm. Ovary 
up to c. 1 mm stipitate, 1-1,5 mm; styles 3 or 4, 
free, 3-4 mm. Fruit (excluding the 1-3 mm 
long gynophore) 1,2-2 cm, containing 1-4 
seeds 6-8 mm. Fig. 38: 2. 

Distributed in eastern South Africa, in the north- 
eastern Cape Province, Natal, Swaziland, and eastern 
Transvaal; extending into Rhodesia, Mozambique and 
southern Tanzania; found in (regularly burnt) grassland, in 
open scrub, forest edges, on sandy and rocky soil; 0-1 700 
m; flowers and fruits can be found throughout the year, but 
mostly in the period Sept. -Oct. 

Transvaal. — 2329 (Pietersburg): Haenertsburg, Pott 
sub TRV 13379. 2330 (Tzaneen): Westfalia Estate, Scheep- 
ers 760. 2530 (Lydenburg): 37 km N.E. of Machadodorp, 
Acocks 16617. 2531 (Komatipoort): Barberton, Galpin 931. 

Swaziland. — 2531 (Komatipoort): Havelock Mine, 
Miller 3075. 2631 (Mbabane): near Mbabane, Compton 

Natal. — 2730 (Vryheid): near Utrecht, Thode A378. 
2831 (Nkandla): near Katazo, Acocks 11795. 2930 (Pieter- 
maritzburg): near Pinetown, Hilliard & Burn 6887. 2931 
(Stanger): near Stanger, Moll 2201. 3030 (Port Shepstone): 
near Highflats, Dyer 4134. 

Cape. — 3029 (Kokstad): Clydesdale, Tyson 2106. 3129 
(Port St. Johns): Mkambati, Codd 9727. 

This is a highly polymorphic species, especially in 
habit, leaf shape, as well as in flower size. The narrow- 
leaved forms described as T. viride are linked up with 
broad-leaved forms by numerous transitional specimens. 

5. Basananthe heterophylla Schinz in 
Verh. Bot. Ver. Prov. Brandenb. 30: 252 
(1888); De Wilde in Blumea 21: 353, fig. 4k, 6 
(1974). Type: S.W. Africa, Amboland, Schinz 
435 (K, iso.!). 

Tryphostemma heterophyllum (Schinz) Engl, in Bot. 
Jahrb. 14: 388 (1891) in obs.; 15: 577 (1893); Pflanzenw. 
Afr. 3, 2: 599 (1921); Hutch. & Pearce in Kew Bull. 1921: 
261 (1921); Harms in Pflanzenfam. ed. 2, 21: 488 (1925); 
Schreiber in F.S.W.A. 89: 2 (1968). 

Annual or biennial to 50 cm, erect, some- 
times ± branched at base, glabrous; tendrils 
absent. Leaves up to 8 X 8 cm, mostly deeply 
3-lobed, sometimes partly simple, the lobes 
oblong-lanceolate, 2-7 x 0,7-2 cm, apex ob- 
tuse to acute, margin up to 1 (-2) mm deep 
serrate-setaceous dentate; leaf-base subcordate 
to subacute; petiole 1-5 cm, shortly winged, 
with gland-teeth. Stipules 3-6 mm; false 



stipules absent. Inflorescences 1- or 2-flowered; 
peduncles 0,2- 1,5 cm; bracts 3-7 mm. Flowers 
glabrous; stipe 2-4 mm; hypanthium 2,5-4 mm 
wide. Sepals 7-15 mm, long-acute, the outer 
with 2 submarginal prominent nerves. Petals 
5-7 mm, (sub)acute. Outer corona-tube 1-1,5 
mm, threads 2, 5-3, 5 mm. Disc 0, 2-0,4 mm. 
Inner corona cup-shaped, 0,5-0, 8 mm. Sta- 
mens 5; filaments (inserted near the base of the 
cup) 4—5 mm, anthers narrow, 2-2,5 mm. 
Ovary 1,5-2 mm; styles 2,5-6 mm, connate for 
1—3,5 mm. Fruit (sub)sessile, 1,5-1, 8 cm, con- 
taining 1-4 seeds c. 7-8 mm. 

Restricted to, but rather widely distributed in the 
northern half of South West Africa, with one collection 
from south-western Botswana. It grows in red or white 
sand, and on sand dunes; 1 000-1 500 m. Flowering and 
fruiting time Dec.-April. 

S.W.A. — 1920 (Tsumkwe): near Tsumkwe, Giess, Watt 
& Snyman 11051. 2115 (Karibib): Karibib, Dinter 6834. 
2116 (Okahandja): Okahandja, Bradfield 323. 

Specimens usually have (deeply) 3-lobed leaves, but 
occasional plants with partly simple leaves have been 

The persistent cotyledons measure 2-2,5 x 1-1,5 cm, 
with a petiole of 1-1,3 cm. 


PassifloraL., Sp. PI. 2; 955 (1753); Gen. PI., ed. 5: 410 (1754); DC., Prodr. 3: 322 (1828); Benth. 
& Hook.f., Gen. PI. 1: 810 (1867); Mast, in Trans. Linn. Soc. Lond. 27: 593 (1871); in FI. Bras. 
13: 531 (1872); Harms in Pflanzenfam. ed. 2, 21: 495 (1925); Killip in Field Mus. Publ. Bot. 19, 1: 
1-613 (1938); Hutch., Gen. FI. PI. 2: 370 (1967); De Wilde, FI. Malesiana 1, 7: 407 (1972). Type: 
P. incarnata L. 

For generic synonyms see Harms, l.c. and Killip, l.c. 

Mostly perennial climbing herbs to large lianas, rarely (not in Africa) shrubs or trees, glabrous 
or hairy, provided with tendrils. Leaves (mostly) alternate, (deeply) lobed or simple, palminerved 
or penninerved, petiolate; margin mostly dentate, often with small gland-teeth; petiole with or 
without glands; blade-glands present or not. Stipules minute to large. Inflorescences sessile or 
peduncled, 1- to many-flowered, with or without a simple tendril; or flowers rarely collected into 
pseudo- racemes; bracts and bracteoles small to large, forming a conspicuous involucre or not. 
Flowers hermaphrodite, 5-merous; hypanthium saucer-shaped to cylindrical. Sepals and petals 
free, often brightly coloured; petals mostly resembling sepals, membranous, sometimes absent. 
Corona extra-staminal, variously shaped, simple or mostly composed of a usually complicated 
outer corona consisting of threads, and flat or plicate inner coronas, with in addition a nectary ring 
or annulus or not. Androgynophore mostly distinct, 3 mm or more. Stamens 5 (-8), free (or in 
some Asian spp. partly connate), in older flowers mostly reflexed; anthers dorsifixed, versatile, 
elliptic to linear. Gynophore absent, or sometimes up to 7 mm. Ovary globose to fusiform; styles 3 
(or 4), free or connate at base; stigmas capitate. Fruit mostly indehiscent, ± baccate, often with 
coriaceous exocarp, globose or ellipsoid, or rarely fusiform, containing many seeds. 

About 370 species, of which c. 350 occur in the Americas, and 20 species in S.E. continental Asia, Indo-Australia and 
the West Pacific. The genus is not indigenous in Africa. Species described from Madagascar and the Mascarene Is. pertain 
to early introductions from America. 

Several species are introduced in South Africa as ornamentals, e.g. P. coerulea L., or for the edible fruits with delicate 
flavour, e.g. P. edulis Sims, P. laurifolia L. and P. quadrangularis L. 

Three species, P.foetida L., P. suberosa L., and P. subpeltata Ortega are locally established weeds in many tropical 
countries; P. edulis Sims is often cultivated and sometimes escapes. These species are marked in the key with an asterisk, 
and treated fully. 

The genus was subdivided by Harms (1925) into 21 sections; Killip (1938) accepts 22 subgenera, and many sections 
and series for the American species. 



Fig. 39. 1, Passiflora coerulea, flowering stem, x 1; a, fruit, x 1 (Robertson s.n.). 



Bracts and bracteoles inconspicuous, filiform or linear, not forming an involucre; flowers small, c. 1,5 cm diam., 

greenish-yellow, without petals I P. suberosa* 

Bracts and bracteoles conspicuous, foliaceous, forming an involucre; flowers much larger, variously coloured, 
provided with petals: 

Involucre bracts finely and deeply divided 2. P.foetida* 

Involucre bracts not divided: 

Leaves not lobed, (sub)pinnately nerved; stem 4-angular, distinctly winged P. quadrangularis 

Leaves usually lobed or partite, palmately nerved; stem not angular or winged: 

Stipules lanceolate or filiform; involucre bracts serrate-denticulate 3. P. edulis* 

Stipules foliaceous; involucre bracts (sub)entire: 

Stipules 1-2 cm long, falcate, remotely dentate; leaves (3-) 5 (-9)-lobed, incisions nearly to the base; corona 

threads with bluish or purplish tinge P. coerulea 

Stipules 1,5-4 cm long, straight, entire; leaves 3-lobed to about the middle; corona threads 

white 4. P. subpeltata* 

1. Passiflora suberosa L., Sp. PI. 2: 958 
(1753); Killip in Field Mus. Nat. Hist. Chicago 
Bot. 19, publ. 407; 88 (1938). Type: ‘Domini- 
ca’ (Hispaniola) (LINN). 

Climber or creeper to 6 m, perennial, 
glabrous or glabrescent or pubescent to various 
degree; stem ± angular, corky when older. 
Leaves entire or usually 3-lobed up to 4 /s the 
blade, subcircular to ovate or oblong in outline, 
3-5-plinerved, membranous or subcoriaceous, 
base rounded or cordate, 4—10 x 4-14 cm, 
lobes triangular to lanceolate, apex acute to 
acuminate; margin entire; petiole 0,5-4 cm, 
with 2 small obconical or wart-like glands at 
about the middle; blade-glands usually absent. 
Stipules linear, 5-8 mm. Inflorescences sessile, 
1- or 2-flowered, with a central tendril, simple, 
3-12 cm; pedicels 1-2 cm, jointed about half- 
way; bracts setaceous, caducous, c. 1 mm. 
Flowers 1-2 cm diam., pale greenish-yellow; 
hypanthium saucer-shaped, 3-5 mm wide; sep- 
als ovate to lanceolate, subobtuse, 5-10 mm; 
petals absent; corona threads in 2 series, 2-6 
mm; operculum (inner corona) plicate, minutely 
fimbriate; disc annular. Androgynophore 2-4 
mm; filaments ± subulate, 2-3 mm; anthers 
1-2 mm; ovary subglobose-ellipsoid, 1-2 mm, 
glabrous; styles 2-3 mm. Fruit a berry, 
(sub)globose, glabrous, 0,8- 1,5 cm diam., 
purple-blackish; seeds several to many, sub- 
ovoid, 3-4 mm long. 

Introduced in many parts of the Old World tropics, up 
to 2 500 m alt; it is sometimes cultivated as an ornamental 
in gardens, while escaped plants can be found on roadsides, 
in disturbed shady places, sandy places near the coast, etc. 

Natal.— 2930 (Pietermaritzburg): near Pieter- 

maritzburg, Acocks 20017. 3030 (Port Shepstone): Ifafa 
River heights, Strey 8312. 

2. Passiflora foetida L., Sp. PI. 2: 959 
(1753); Killip in Field Mus. Nat. Hist. Chicago 

Bot. 19: 474 (1938). Type: South America, 
(perhaps Lesser Antilles; fide Killip) (LINN). 

Climber or creeper to 4 m, annual or 
biennial, subglabrous or whitish or yellowish- 
brown hairy to various degree; ill-odoured; 
stem terete. Leaves entire, or usually 3 (-5)- 
lobed up to half-way, suborbicular to ovate in 
outline, 3-5-subplinerved, usually membran- 
ous, base cordate, 3-10 x 3-10 cm; lobes up to 
4 cm, usually acute-acuminate, margin (sub)en- 
tire with coarser gland-tipped hairs; petiole 1-6 
cm; stipules subreniform, 0,5-1 cm, deeply 
cleft into filiform gland-tipped processes; 
glands absent, except gland-tipped stronger 
hairs or processes on petiole, stipules, bracts, 
etc. Inflorescences sessile, 1 (-2)-flowered, the 
straight peduncle 2-6 cm inserted beside a 
simple tendril 5-15 cm; bracts and bracteoles 
(1-) 2-4 cm, deeply 2-4 times pinnatisect, with 
filiform gland-tipped segments, forming an in- 
volucre just below, and enveloping the flower. 
Flowers c. 3-5 cm diam., pale pinkish or lilac, 
rarely white; hypanthium short, saucer-shaped; 
sepals ovate-oblong to lanceolate, c. 1,5-2 cm, 
awned 2-4 mm dorsally just below the apex; 
petals oblong to lanceolate or ± spathulate, 
slightly shorter than the sepals; corona consist- 
ing of 2 outer series of threads, c. 1 cm long, 
several inner series of capillary threads 1-2 
mm; operculum membranous, ± erect, denticu- 
late; disc conspicuous, annular. An- 
drogynophore 4-6 mm; filaments flattened, c. 
5-6 mm; anthers 3-5 mm; ovary globose to 
ellipsoid, c. 2-3,5 mm, usually glabrous; styles 
4—5 mm. Fruit a rather dry berry, (sub)globose, 
usually glabrous, 1,5-3 cm diam., yellowish to 
orange, ± enveloped by persistent involucre; 
seeds many, subovoid to ± cuneiform, c. 4-5 
mm long, obscurely 3-dentate at apex. 

A common alien in parts of the subtropics and all 
tropical areas, up to 2 500 m altitude. It is found on coastal 



sands, in waste places, in ruderal and disturbed vegetation, 
and roadsides. It is sometimes cultivated and often escapes. 
Killip (l.c.) recognized many varieties, but most African 
material can be referred to var. hispida (DC.) Killip ex 
Gleason, a variety with glabrous ovary, and usually 3-lobed 
leaves, the whole plant with a rather hispid tomentum. 

Natal. — 2930 (Pietermaritzburg): Merebank, Baijnath 
180; Isipingo Flats, Ward 6244. 

3. Passiflora edulis Sims in Bot. Mag. 45: 
1. 1989 (1818); Killip in Field Mus. Nat. Hist. 
Chicago Bot. 19: 393 (1938). Type: cult, in 
Europe, probably originally from Brazil (fide 
Killip, l.c.). 

Climber to c. 15 m, perennial, glabrous 
throughout (except ovary); stem sometimes ± 
angular. Leaves 3-lobed up to % the blade, 
rarely without lobes, suborbicular to broadly 
ovate in outline, 5-11 x 6-12 cm, 3-plinerved, 
subcoriaceous, base acute to cordate, lobes 
elliptic to oblong, up to 8 cm, apex acute, 
shortly acuminate; margin serrate; petiole 1-4 
cm; glands on petiole 2, wart-like, situated at 
transition of or up to 0,5 cm below the blade; 
blade-glands absent. Stipules lanceolate-linear, 
c. 1 cm. Inflorescences 1 -flowered, the straight 
peduncle (pedicel) 3-6 cm, inserted beside a 
simple tendril 5-20 cm; bracts and bracteoles 
ovate, acute, 1,5-2 cm, glandular-serrate, form- 
ing an involucre. Flowers 4—7 cm diam., white, 
corona threads purplish towards base; hypan- 
thium cup-shaped, c. 1 x 1—1,5 cm; sepals 
oblong, 2-3 cm, comiculate; petals oblong, 
1-2,5 cm, obtuse; corona composed of several 
series of threads, the outer 2 series 0,5-2, 5 cm, 
the inner ones much shorter; operculum mem- 
branous, incurved, crenulate-fimbriate; disc 
(limen) cupuliform, entire or crenulate. An- 
drogynophore 6-8 mm, thickened towards 
base; filaments sub-subulate, 6-8 mm; anthers 
8-10 mm; ovary subglobose to ellipsoid, 3-5 
mm, glabrous or shortly pubescent; styles 
10-12 mm. Fruit berry-like, with coriaceous- 
leathery pericarp, globose to ellipsoid, exclud- 
ing the 0,5- 1,5 cm long gynophore, 4-5 cm 
diam., glabrous, purplish or yellow; seeds 
many, ellipsoid, 5-6 mm. 

This species is cultivated for its edible, well-flavoured 
fruit. Sometimes escaped in waste gardens, disturbed places 
and forest fringes; in the tropics up to 2 500 m alt. 

Transvaal. — 2230 (Messina): near Lake Funduzi, Story 
4847. 2527 (Rustenburg): Jacksonstuin, Van 

Vuuren 196. 2530 (Lydenburg): Vertroosting Nature Re- 
serve, Muller 2492 , 2531 (Komatipoort): Ida Doyer Nature 
Reserve, Muller 2119. 

Natal. — 2831 (Nkandla): Port Dumford, Venter 1882. 

4. Passiflora subpeltata Ortega, Nov. 
Rar. PI. Hort. Madrit. 6: 78(1798); Killip in Field 
Mus. Nat. Hist. Chicago Bot. 19: 436 (1938). 
Type: Mexico (specimen in Herb. Madrid, fide 

Herbaceous creeper or climber to 5 m, 
perennial, essentially glabrous throughout; stem 
terete. Leaves 3-lobed to about half-way, sub- 
orbicular in outline, 4-10 x 4—11 cm, (3-) 
5-plinerved, herbaceous, base rounded, truncate 
or cordate, subpeltate; lobes elliptic to oblong, 
up to 5 cm, apex obtuse or acutish, c. 1 mm 
mucronate; margin entire except for a few 
gland-teeth in or near the lobe-sinuses; petiole 
3-6 cm; glands on petiole 2-5, scattered or ± 
paired at about the middle, slender, up to 1 mm 
long; blade-glands absent. Stipules ovate- 
oblong, straight, 1,5-4 cm long, entire or with a 
few minute gland-teeth, apex mucronulate. 
Inflorescences 1 -flowered, the straight peduncle 

3- 6 cm, inserted besides a simple tendril 4—12 
cm; bracts and bracteoles (broadly) ovate 1-1,5 
cm, acute, entire or with a few minute gland- 
teeth at base, forming an involucre. Flowers 

4— 5,5 cm diam., white; hypanthium broadly 
cup-shaped, 7-10 mm wide; sepals oblong, 
2-2,5 cm, obtuse, with a subapical horn 0,5-1 
cm; petals oblong, 1,5-2 cm, acutish; corona 
composed of 4 or 5 series of threads, those of 
the outer 2 series 1 (-1,5) cm long, those of 
inner series 2-6 mm; operculum subplicate, 
fimbriate-laciniate for about half or less, and 
with a fringe of inward curved dentiform pro- 
cesses; disc annular; limen with lobulate edge, 
erect or ± reflexed at the top, closely surround- 
ing androgynophore. Androgynophore 10-12 
mm; filaments 5-6 mm, dilated; anthers 5-7 
mm; ovary ellipsoid, c. 4 mm, glabrous; styles 
8-10 mm. Fruit ± leathery, ellipsoid or sub- 
globose, excluding the 1,5-2 cm long 
gynophore 3, 5-4, 5 cm long, greenish turning 
yellow; seeds many, ellipsoid, 4-5 mm. 

Originating from tropical America and cultivated as an 
ornamental; sometimes profusely escaped, e.g. in aban- 
doned plantations, on forest edges, etc. ; up to 1 000 m, in 
the tropics up to 2 500 m. 

Transvaal. — 2530 (Lydenburg): Lowveld Botanic Gar- 
den, Buitendag 615. 

Natal. — 2930 (Pietermaritzburg): Nagle Dam, 1 Veils 
1354. 3030 (Port Shepstone): Umgai, Strey 10583. 

Killip, in 1947, named the identical material in the East 
African Herbarium as the related and similar species P. 
eichlerana Mast. (1872). This species, according to Killip 
(1938), differs from true P. subpeltata mainly by the 
operculum, which in P. eichlerana is laciniate for about 



half or more, and by the presence of inward curved 
dentiform processes; in P. subpeltata the operculum is 
fringed only at the margin, with the entire portion much 
longer than the teeth or fringes. All the abundant material 
now available from East Africa, as well as similar material 
from India, Malesia, Australia, Hawaii, etc., is doubtlessly 
conspecific. The operculum in all these alien specimens is 

dissected up to about half-way or less, thus rendering the 
distinction between P. eichlerana and P. subpeltata for 
these specimens difficult. As both species are related in their 
original country, the introduced plants are possibly a deviat- 
ing form of one or the other, or a hybrid. As P. subpeltata is 
the older name, I have chosen to accept it over P. 


by D. J. B. Killick 

Herbaceous climbers, subherbaceous shrublets or acaulescent herbs; leaves alternate, simple, 
lobed or not lobed, sometimes all radical, crenate or serrate; stipules present or 0. Plants 
monoecious. Inflorescence a raceme or flowers 1-several, axillary. Sepals 3-5, usually linear, 
sometimes adnate to corolla-tube. Corolla-tube campanulate; lobes 3-4, about as long as the tube 
or shorter. Stamens 3-5, inserted at base of corolla-tube or in throat; anthers 2-thecous, sometimes 
cohering. Glands inserted at base of corolla-tube. Ovary superior, 1-locular, with few to many 
ovules on 3-5 parietal placentas; style 3-5-lobed with the lobes sometimes 2-fid. Fruit a capsule, 
enclosed in a perianth corolla. Seed subglobose, pitted or tubercled; embryo straight; cotyledons 
flat; endosperm present. 

Genera 3, species 3, occurring in South Africa and Swaziland. Achariaceae was included by Bentham & Hooker f. 
(1867) in Passifloraceae, but differs in the sympetalous campanulate corolla and, in two genera, the stamens which are 
adnate to the corolla-tube. According to Hutchinson (1967) the habit of Ceratiosicyos recalls that of some Cucurbitaceae, 

hence Achariaceae represents something of a link between Passifloraceae and Cucurbitaceae. 

Acaulescent herbs with radical leaves 3. Guthriea 

Plants not acaulescent; 

Erect shrublets; male flowers axillary, 1-several; sepals present in female flowers; fruit 

ellipsoid, up to 1 cm long 2. Acharia 

Herbaceous climbers; male flowers racemose; sepals absent in female flowers; fruit 

siliquiform, up to 8 cm long 1. Ceratiosicyos 


Ceratiosicyos Nees in Eckl. & Zeyh., Enum. 281 (1836); Harv., Gen. S. Afr. PI. 107 (1838); in 
F.C. 2: 501 (1862); Benth. & Hook, f., Gen. PI. 1, 3: 814 (1867); Phill., Gen. ed. 2; 519 (1951). 
Type species: C. ecklonii Nees. 

Herbaceous climber. Leaves alternate, simple, petiolate, exstipulate. Male flowers racemose. 
Sepals 4 or 5. Corolla-tube campanulate; lobes 4 or 5. Stamens 5, inserted at base of corolla-tube, 
alternating with an equal number of glands adnate to corolla-tube; filaments free. Ovary 0. Female 
flowers solitary. Calyx absent. Corolla as in male flowers. Glands 5. Ovary shortly stalked, with 
several ovules on 4 or 5 parietal placentas; styles 4 or 5, 2-lobed at apex. Fruit an elongated ribbed 
capsule, several seeded. Seeds with tuberculate testa; embryo straight; cotyledons ovate, distinctly 
veined; endosperm copious, fleshy. 

One species found in the Transvaal, Swaziland, Natal and the Cape. The generic name is compounded from the Greek 
for pod or siiiqua and cucumber in reference to the nature of the fruits. 



Fig. 40. — 1, Ceratiosicyos laevis, flowering and fruiting stem, X 1; a, male flower, X 5; b, corolla of male flower opened 
out, X 6; c, upper part of stamens, front and side views, x 5; d, fruit, X 1; e, female flower, x 8; f, corolla of female 
flower opened out; g, pistil, x 8; h, stigma, X 10; i, portion of ovary opened out to show attachment of ovules, x 20 
(mostly after Harvey in Ann. Nat. Hist. ser. 1, 3: 1. 10, 1839). 



Ceratiosicyos laevis (Thunb.) A. Meeuse 
in Bothalia 8: 20 (1962). Type: Cape, Thunberg 
s.n. (UPS, holo. !; PRE, photo.). 

Bryonia laevis Thunb., Prodr. 1: 13 (1794). 

Ceratiosicyos ecklonii Nees in Eckl. & Zeyh., Enum. 281 
(1836); Harv., Gen. S. Afr. PI. 107 (1838); Am. & Harv. in 
Ann. Nat. Hist. ser. 1, 3: 421, 1. 10 (1839); Harv. in F.C. 2: 
501 (1862); Marloth, FI. S. Afr., 2,2: 200, 1. 131 (1925). 
Type: Cape, “in nemoribus ad Krakakamma, Adow & 
Olifantshoek (Uitenhage), turn in Albany & Kafferland”, 
Ecklon 1797 (BOL!; G!; K!; L!; S!). 

Herbaceous climber, glabrous. Leaf- 
blade palmately 5-7-lobed with lobes acumi- 
nate and margins serrate, 3-14 cm long, 3-12 
cm wide; petiole 2-14 cm long. Male flowers 
yellowish-green. Sepals filiform-linear, 2,5-3 
mm long, 0,2 mm wide, slightly swollen at 
apex, spreading. Corolla-tube 3,5-5 mm long; 
lobes slightly shorter than the tube, ovate- 
oblong, ciliate, cohering together by the cilia 
giving the corolla an urceolate appearance. 
Glands linear or obovate, 2 mm long, fleshy. 
Stamens with linear filaments, 5-6 mm long, 
dilated upwards; anthers oblong, c. 3 mm long, 
cohering, reflexed at apex. Female flowers 
yellowish-green. Corolla-tube 2-3,5 mm long; 
lobes 2-3 mm long. Glands narrowly ovate, up 
to 1,5 mm long, fleshy. Ovary stipitate, linear- 
elongate, 3-4 mm long; styles short, 2-lobed 
with recurved lobes. Fruit elongated, cylindri- 
cal up to 8 cm long, 0,8 cm wide, tapered at 


both ends. Seeds subglobose to oblong, 3-4 
mm diam., tuberculate. Fig. 40. 

A forest climber distributed from Wilderness in the 
south-eastern Cape, into Natal and Swaziland, and north- 
wards as far as Duiwelskloof in the northern Transvaal. 

Transvaal. — 2330 (Tzaneen): Westfalia Estate, 

Duiwelskloof, Scheepers 397. 2430 (Pilgrims Rest): 1,6 km 
from Graskop to Kowyns Pass, Jordaan 100; The Downs, 
Junod 4197. 

Swaziland. — 2631 (Mbabane): Duiker Bush, Compton 

Natal. — 2731 (Louwsburg): Wendelane Spruit, Non- 
goma, Gerstner 4417. 2829 (Harrismith): Draycott Hill, 
Acocks 11436. 2830 (Dundee): 20 km from York on 
Rietvlei Road, Marais 350. 2831 (Nkandla): Ngoye, 
Ubisana Valley, Venter 1313. 2832 (Mtubatuba): Hluhluwe 
Game Reserve, Hitchins & Ward 2. 2929 (Underberg): 
Deepdale, near Umkomaas River, Strey 4815. 2930 
(Pietermaritzburg): “Ehlatini”, Karkloof, Moll 3468. 2931 
(Stanger): Umgeni River Mouth, Schlechter 2845 (K). 3030 
(Port Shepstone): Uvongo, Strey 9598; Dumisa, Rudatis 
470 (K). 

Cape. — 3029 (Kokstad): Mount Currie, Tyson 472. 3126 
(Queenstown): N’Zebanya Mountain, near Queenstown, 
Galpin 1932. 3128 (Umtata): 23 km from Umtata on road to 
Ngqeleni, Story 1327. 3226 (Fort Beaufort): Katberg, 
Hutton s.n. (K). 3227 (Stutterheim): Hogsback, Acocks 
11018. 3228 (Butterworth): Kentani, Pegler 903. 3322 
(Albert): Wilderness, Van Niekerk 213. 3323 (Willow- 
more): Stinkhoutkloof, Knysna, Forest Officer s.n. 3325 
(Port Elizabeth): “Vista”, ± 5 km from Paterson on road to 
Mimosa, Retief 18. 3326 (Grahamstown): Alexandria 
Forest, Johnson 1059. 

A readily distinguishable herbaceous climber with 
5-7-lobed leaves and elongated, ribbed capsular fruit up to 
8 cm long. 


Acharia Thunb., Prodr. 1: 14, t. s.n. (1794); FI. Cap. ed. Schult. 37 (1823); Harv., Gen. S. Afr. PI. 
409 (1838); Am. & Harv. in Ann. Nat. Hist. ser. 1,3: 420, t.9 (1839); Harv. in F.C. 2: 501 (1762); 
Benth. & Hook.f., Gen. PI. 1,3: 814 (1867); Phill., Gen. ed. 2: 520 (1951). Type species: A. 
tragodes Thunb. 

Shrubby herb. Leaves alternate, simple petiolate, stipulate. Flowers axillary, 1-several, 
cemuous. Male flowers: Sepals 3 or 4. Corolla pubescent; tube campanulate; lobes 3 or 4, slightly 
winged or keeled. Glands 3 or 4, at base of corolla-tube, ciliate. Stamens 3 or 4, free; anthers 
oblong with cells widely separated. Ovary 0. Female flowers: Sepals 3 or 4, larger than in male, 
pubescent. Corolla larger than in male, pubescent; tube campanulate, keeled; lobes ovate. Glands 
3 or 4, at the base of the corolla-tube. Ovary subsessile or stalked, with 3-5 ovules on 3-5 parietal 
placentas, densely pubescent; style 3-5-lobed. Fruit a capsule, enclosed in persistent corolla, 
3-5-valved, terminated by the persistent style. Seed 1, narrowly winged (arillate) on one side. 

An endemic, monotypic genus extending from the Uitenhage District in the Cape into Natal. The genus was named in 
honour of Erik Acharius (1757-1819), pupil of Linnaeus, lichenologist and physician of Vadstena in Sweden. 

Acharia tragodes Thunb., Prodr. 1: 14, t. s.n. (1794); FI. Cap. ed. Schult. 37 (1823); Harv., 
Gen. S. Afr. PI. 409 (1838); Am. & Harv. in Ann. Nat. Hist. ser. 1,3: 420, t.9 (1839); Harv. in F.C. 
2: 502 (1862); Martin & Noel, FI. Albany & Bathurst 77 (1960). Type: Thunberg s.n. (UPS, 



Fig. 41. — 1, Acharia tragodes, flowering stems, X 1; a, male flower, x 5; b, view of male flower from above, x 4; c, 
sepals, X 3; d, corolla tube opened out, x 4; e, basal glands, X 6; f, glandular anther, x 10; g, female flower, x 4; h, 
pistil with basal glands, X 10; i, stigmas, X 10; j, stalked fruit, X 3; k, dehiscing fruit, X 3; 1, fruit valve with seeds, X 
3; m, seed, X 4 (mostly after Harvey in Ann. Nat. Hist. ser. 1, 3: t.9, 1839). 



Perennial shrubby herb up to 40 cm high. 
Leaf -blade deeply palmately-lobed, 1,5-6, 3 cm 
long, 1,5-6 cm wide, subchartaceous, pubes- 
cent; petiole 1,5-3 cm long, pubescent. Flow- 
ers 1 or more (of one or both sexes) together, 
shortly pedicellate. Male flowers: Sepals linear 
to narrowly elliptic, 1,8-3 mm long, 0,5-0,75 
mm wide, somewhat recurved, pubescent. 
Corolla-tube 3-4 mm long; lobes triangular, 
2-2,2 mm long. Glands ovate, 1 mm long, 0,75 
mm wide. Stamens with filaments 0,5-1, 2 mm 
long; anthers 1-1,5 mm long, glandular. 
Female flowers: Sepals elliptic, 2-3,5 mm long, 
1,5 mm wide. Corolla-tube 4—5,5 mm long, 
lobes ovate, 3 mm long, 3-4 mm wide. Glands 
ovate, c. 1 mm long, pubescent. Ovary subses- 
sile or with stalk 1 mm long, ellipsoid, 2-2,5 
mm long, 1,5-2 mm wide; style terete, 2 mm 

long, lobes 1 mm long, with flabelliform stig- 
mas. Fruit ellipsoid, 8-10 mm long, 5-8 mm 
wide, pubescent. Seed subglobose, 3 mm 
diam., reticulate. Fig. 41. 

A rather rare species occurring in scrub and woods 
from the Uitenhage District in the Cape coastwise to Durban 
in Natal. 

Natal. — 2931 (Stanger): Port Natal, Drege s.n. (G). 

Cape. — 3228 (Butterworth): Kentani, Pegler 757; near 
Kei River Mouth, Flanagan 50; Collywobbles, Idutywa, 
Van Breda 876. 3325 (Port Elizabeth): Van Stadensberg, 
Zeyher 4654 (P;S); Elandsrivier, Ecklon & Zeyher 105,2; 
Wall 17 (S). 3326 (Grahamstown): Hopewell, Acocks 

A perennial, shrubby herb growing up to 40 cm high 
with deeply palmately-lobed leaves and ellipsoid fruit up to 
1 cm long. The specific epithet has been variously spelled as 
tragodes, tragoides and tragioides, but there is no reason 
why there should be any departure from the original 
spelling, tragodes. 

5376 3. GUTHRIEA 

Guthriea H. Bol. in Hook. Icon. PI. 1. 1 16 1 (1876); Marloth, FI. S. Afr. 2,2: 200 (1925); Phill. 
Gen. ed. 2: 520 (1951). Type species: G. capensis H. Bol. 

An acaulescent herb. Leaves rosulate, alternate, simple, petiolate. Flowers axillary, solitary, 
pedicelled. Male flowers: Sepals 5, adnate to corolla for almost whole of their length, shortly 
exceeding corolla. Corolla campanulate; lobes 5, reflexed. Glands 5, fleshy, inserted at base of 
corolla-tube. Stamens 5, inserted in throat of corolla; filaments slightly longer than anthers, 
flattened; anthers with cells attached to a broad connective. Female flowers: Sepals 4 or 5, adnate 
to corolla for half their length, about half as long as corolla. Corolla-tube campanulate; lobes 4 or 
5. Glands 4 or 5, fleshy, inserted at base of corolla-tube. Ovary sessile, with 10-15 ovules on 4 or 
5 parietal placentas; style 4- or 5-lobed. Fruit a 4- or 5-valved capsule enclosed in the corolla. 
Seeds several, arillate; embryo straight. 

A genus of one species occurring in the eastern Cape, Natal and Lesotho. The genus was named in honour of Francis 
Guthrie, Professor of Mathematics, South African College, Cape Town. 

Guthriea capensis H. Bol. in Hook. Icon. 
PI. 1. 1 161 (1876); Marloth, FI. S. Afr. 2,2: 200 
(1925). Type: Cape, “in saxosis in monte 
Gnadouw-Sneeuwbergen in ditione Graaff 
Reinet”, et “in monte Oudeberg prope Graaff 
Reinet in locis graminosis”, Bolus 818 (BOL, 
holo. !; K!; PRE!; SAM!; Z!). 

An acaulescent herb with a sub-horizontal 
rhizome and thick, fleshy roots. Leaf-blade 
broadly ovate, occasionally broadly elliptic, 
4—7 cm long, 3, 5-6, 5 cm wide, apex obtuse to 
round, base cordate, margins crenate, digitately 
nerved, discolorous, dark green and shiny 
above, paler below, bullate, glabrous; petiole 
3-12 cm long. Flowers with pedicels 2-8 cm 
long. Male flowers: Sepals linear, 1,5 cm long, 
projecting beyond corolla, c. 2,5 mm. Corolla 

1,3-1, 5 cm long, 1,2 cm wide at mouth, pubes- 
cent inside in lower half; lobes ovate, 4 mm 
long, 3 mm wide. Glands ovate, 2 mm long, 1,5 
mm wide, thick, fleshy, crearri. Stamens with 
filaments 3,5 mm long, anthers 2-2,5 mm long, 
glandular. Female flowers: Sepals linear, c. 6 
mm long, free for 2-3 mm. Corolla-tube 1,3- 
1,5 cm long, 1,2-1, 4 cm wide at mouth; lobes 
transversely ovate, 1-3 mm long, 6 mm wide, 
erect. Glands ovate to obovate, 2,5 mm long, 2 
mm wide, cream. Ovary ellipsoid, 8 mm long, 
4 mm wide, brown gland-dotted; style subter- 
ete, 4-5 mm long; lobes simple, 1 mm long, 
spreading. Fruit triangular-ellipsoid, 1,2 cm 
long, 1 cm wide. Seeds ellipsoid, 3,5 mm long, 
2 mm wide, minutely tuberculate, pubescent, 
black. Fig. 42. 



Fig. 42. — 1, Guthriea capensis, flowering plant, x 1; a, male flower opened out, x 1 16; b, female flower opened out, x 
116; c, transverse section of ovary, X 2; d, fruit with persistent style and corolla, X 116; e, seed, x 5 (Mostly after Fitch 
in Hook. Icon. PI. 1. 1 161, 1876). 



Herb occurring in grassland on mountains in the 
eastern Cape and Natal from the Sneeuwbergen to Mont aux 
Sources and in Lesotho. 

Natal. — 2828 (Bethlehem): Sentinel area, Mont aux 
Sources, Killick & Marais 2199; Schweickerdt 707 ; summit 
of Mont aux Sources, Flanagan 2103. 2929 (Underberg): 
Giants Castle, Symons 345. 

Lesotho. — 2928 (Marakabies): Mamalope, Jacot Guil- 
larmod 766. 

Cape. — 3027 (Lady Grey): Ben McDhui, Wittebergen, 
Galpin 6630. 3028 (Matatiele): Naude’s Nek, Hilliard 5212 
(NBG). 3124 (Hanover): Compassberg, Acocks 16424. 

3224 (Graaff Reinet): Oudeberg, Bolus 818; 1044 (K); 
“Gnadouwsberg”, Bolus 818. 

An acaulescent herb with radical, rosulate, bullate 
leaves dark green and shiny above and paler below, with 
flowers mostly hidden under the leaves. Bolus lumped his 
type material of G. capensis from Gnadouw Mountain 
(probably Nadouwsberg, 40 km E. of Graaff Reinet) and 
Oudeberg (probably Ouberg 20 km N.W. of Graaff Reinet) 
under one number. Bolus 818. The specimen in Bolus 
Herbarium from Gnadouw Mountain with notes by Bolus in 
his own handwriting is presumably the holotype rather than 
the one from Oudeberg marked ‘typus auctoris’, but not in 
Bolus’s hand. 


by L. E. Codd 

Herbs, rarely woody, mostly clothed with rough hairs. Leaves alternate or opposite, entire or 
variously divided, exstipulate. Inflorescence a raceme or cyme, rarely a head or flowers solitary; 
bracteoles present. Flowers bisexual, regular. Calyx-tube adnate to the ovary; lobes 4-5, imbricate 
or contorted, persistent, often enlarged in fruit. Petals 4—5, free, inserted in the throat of the 
calyx-tube, induplicate-valvate. Nectary scales petaloid, cucullate, alternating with the petals, 
appendaged or scales absent. Stamens many, rarely few, inserted with the petals, often collected in 
bundles opposite the petals; filaments sometimes beaded; anthers often didymous, 2-thecous, 
opening by longitudinal slits; staminodes present or absent. Ovary inferior or nearly completely so, 

1- 3-chambered, with 1-many pendulous ovules from the apex of the ovary chambers or on parietal 
placentas; style entire or 2-3-fid; stigma simple or capitate. Fruit a capsule, 1-3-locular, 

2- 5-valved. Seed often minute; embryo straight; endosperm present or absent. 

Genera 15, species about 230; 14 of the genera and all but two of the species are natives of tropical or South America, 
with the remaining genus, Kissenia, in Africa. 


Kissenia R. Br. exEndl., Gen. PI. Suppl. 2: 76 (1842) (err. Fissenia ); T. Anders, in J. Linn. Soc. 
(Bot.) 5, Suppl. 1:43 (1861); Harv. in F.C. 2: 503 (1862); Gen. ed. 2: 1 19 (1868); Mast, in F.T.A. 
2: 501 (1871); Gilg in Pflanzenfam. 3, 6a: 114 (1894); Phill. , Gen. ed. 2: 521; (1951); Dandy in 
Kew Bull. 20: 451 (1966). 

Subshrub, scabrous, covered with brittle hairs armed with minute recurved barbs. Leaves 
alternate, petiolate, slightly or deeply lobed. Inflorescence a scorpioid, bracteate cyme. Calyx-tube 
obconic, 10-ribbed, densely villous; lobes 5, equal, longer than the tube, oblong-spathulate, 
enlarged in fruit. Petals shorter than the calyx-tube, concave, hairy without. Nectary scales 5, 
petaloid. Stamens many, some of the outer stamens with a hairy scale at the base and with or 
without anthers; filaments linear, all or mostly beaded. Ovary inferior, 2-3-chambered, with 1 or 
rarely 2 apical pendulous ovules in each chamber; the portion of the ovary projecting above the 
calyx-tube flattened and densely villous; style simple, somewhat angled; stigma simple. Fruit 
woody, ribbed, 2-3-locular, 1-3-seeded, indehiscent, crowned with the enlarged calyx-lobes, 
densely villous. Seed compressed; cotyledons flat, thick, fleshy, oblong; endosperm 0. 



A genus of 2 closely related species with a disjunct distribution, one in southern Arabia, Somalia and Ethiopia and the 
other in the western parts of Southern Africa. 

The name is derived from a place in southern Arabia now known as Kishin or Qishn (Dandy, 1966). Fissenia is 
considered to be an unintentional orthographic error on the part of Endlicher for Kissenia, the ms. name provided for it by 
Robert Brown. 

Kissenia capensis End!., Gen. PI. Suppl. 
2: 76 (1842) (err. Fissenia)-, Harv., Thes. Cap. 
1: 61 (1859) (err. Fissenia)-, Dandy in Kew 
Bull. 1926: 111 (1926); in Kew Bull. 20: 451 
(1966); Roessler in F.S.W.A. 93: 1 (1968). 
Type: Namaqualand, between Verleptpram and 
Orange River Mouth, Drege. 

Cnidone mentzelioides E. Mey. ex Drege, Cat. PI. Exsicc. 
Afr. Austr. 27 (1838), nont. nud.; Zwei Pfl. Doc. 93 (1843), 
nom. nud. 

Kissenia mentzelioides R. Br. ex Harv., Thes. Cap. 1: 
sub. t.98 (1859) (err. Fissenia), nom. syn. K. spathulata R. 
Br. ex T. Anders, in J. Linn. Soc. (Bot.) 5, Suppl.: 43 
(1861); Harv. in F.C. 2: 503 (1862); Mast, in F.T.A. 2: 501 
(1871); Marloth, FI. S. Afr. 2,2: 200, t.68, f. 132 (1925); 
nom. illegit. 

Pallid, scabrid subshrub 25-120 cm tall; 
bark whitish. Leaves ovate to ovate-oblong, 
6-9 cm long, 4—7 cm broad, markedly scabrid, 
usually 3-5-lobed; base truncate to subcordate, 
apex obtuse; margin irregularly dentate; petiole 
2-5 cm long. Flowers in terminal, scorpioid 
cymes; bracts ovate-lanceolate, 5-9 mm long, 
persistent. Calyx-tube densely hirsute; lobes 
pale green, turning yellow, oblong, 17-20 mm 
long, 4 mm broad, finely hispid on both sur- 
faces. Petals white, spathulate, concave, 9-12 

mm long, 5 mm broad, pubescent without. 
Nectary scales usually 5, sometimes fewer or 
with one smaller than the rest, strap-shaped, 
cucullate, 4—5 mm long, 1 mm broad, recurved 
at the apex, hairy within. Stamens, ovary, fruit 
and seed, as for generic description. Fig. 43. 

Often locally common in sandy soil between boulders, 
on river banks and dry water courses in north-western Cape 
Province and the western half of S.W. Africa. 

S.W.A. — 1812 (Sanitatas): Anabib, De Winter & Leist- 
ner 5742. 2014 (Welwitschia): Welwitschia, Galpin & 
Pearson 7631. 2015 (Otjihorongo): 65,6 km W. of Outjo, 
De Winter 3062 A. 2114 (Uis:) near Brandberg, Rodin 
2747. 2115 (Karibib): Usakos, Marloth 1226. 2215 (Trek- 
kopje): Namib Desert Park, Jensen 90. 2315 (Rostock): 
near Gobabeb, Koch A 21. 2316 (Nauchas): Farm Ubib, 
Merxmiiller & Giess 28116. 2416 (Maltahohe): Buellsport, 
Strey2150. 2518 (Tses): Karub, Wiss972. 2617 (Bethanie): 
Seeheim, De Winter & Giess 6421a. 2618 (Keetmanshoop): 
Keetmanshoop, Dinter 3556. 2718 (Grunau): Klein Karas, 
Oertendahl 90a. 2818 (Warmbad); near Warmbad, Ver- 
doorn & Dyer 1776. 2819 (Ariamsvlei): Ham River, Galpin 

Cape. — 2817 (Vioolsdrift): Richtersveld, Hardy 1665. 
2818 (Warmbad): Wolveton, Schlechter 11439. 2820 
(Kakamas): Keimoes, Marloth 14029. 2822 (Glen Lyon): 
Langeberg, Acocks 495. 2917 (Springbok): 48 km N. of 
Steinkopf, Hartmann 1587. 2919 (Pofadder): near Pella, 
Pearson 3544. 2922 (Prieska): 21 km E. of Draghoender, 
Codd 1228. 2923 (Douglas): Kameelfontein, Bryant J1 14. 


by O. M. Hilliard* 

Herbs or undershrubs, mostly fleshy, generally erect, sometimes creeping or climbing by 
means of adventitious roots, or acaulescent with a rhizome or tuber. Leaves alternate, rarely 
subradical, petiolate, generally asymmetrical, digitately nerved, margins entire, toothed, lobed or 
dissected. Stipules 2, conspicuous, persistent or caducous. Inflorescences axillary, cymose, 
bracteate. Flowers monoecious, actinomorphic or zygomorphic, showy. Male flowers: tepals 
usually 2 or 4, rarely many or 0, petaloid. Stamens numerous, filaments free or connate; anthers 
2-thecous, dehiscing laterally. Ovary rudimentary or 0. Female flowers: tepals 2-5, rarely many, 
petaloid. Staminodes 0 or rarely represented by glands. Ovary inferior or rarely half-inferior, 2-4-, 
rarely 1-, celled, usually 3-celled and 3-winged or angled, ovules numerous on projecting simple or 
lobed axile placentae. Styles 2-5, free or connate at the base, usually 2-fid, the stigmas often 

*Department of Botany, University of Natal, Pietermaritzburg. All type specimens cited were seen by the author unless 
otherwise stated. 



twisted, papillose. Fruit a capsule, rarely a berry. Seeds numerous, minute, testa reticulate, 
endosperm scanty or 0, embryo straight. 

A family of 5 genera and about 900 species, most of them in the genus Begonia. Mainly natives of the warm, wet parts 
of America and Asia, relatively few in Africa and represented here only by the genus Begonia. 


BegoniaT., Sp. PI. 1: 1056 (1753); Gen. PI. ed. 5: 475 (1754); Klotzsch in Abh. Ak. Wiss. Berlin 
1854: 125 (1855), reimp. Beg. Gattung Art. (1855); Sond. in F.C. 2: 480 (1862); A. DC. in DC., 
Prodr. 15: 266 (1864); Benth. & Hook.f., Gen. PI. 1: 841 (1867); Hook.f. in F.T.A. 2: 569 (1871); 
Warb. in Pflanzenfam. 3,6a: 121 (1894); Engl., Pflanzenw. Afr. 3,3: 612 (1921); in Pflanzenfam. 
ed. 2, 21: 548 (1925); Phill. , Gen. ed. 2: 522 (1951); Irmscher in Bot. Jahrb. 81: 106 (1961). 

Caulescent or acaulescent herbs or undershrubs with watery stems and leaves, generally erect, 
sometimes creeping or climbing, sometimes with rhizomes or tubers. Leaves alternate, rarely 
subradical, petiolate, usually asymmetrical, entire, lobed or partite, irregularly toothed, green or 
sometimes richly coloured or spotted. Male flowers : tepals 2 or 4, rarely many or 0, petaloid. 
Stamens numerous, filaments free or connate at the base. Female flowers: tepals often 5 or 6, 
sometimes 4, petaloid. Ovary’ usually 3-, rarely 2-, 4- or 5- celled; ovules numerous, placentae 
axile, projecting, simple or lobed. Styles usually as many as the loculi, free or connate at the base, 

2- fid, the stigmas entire or branched, often twisted, papillose. Fruit usually a capsule, 3-winged or 

3- angled, rarely terete or 4-angled, or a berry. Seeds numerous, minute, endosperm 0. 

About 900 species described, mainly from tropical America and Asia. Perhaps only 5 native in South Africa. 

Placentae bilamellate (they appear bifurcate in transverse section): 

Stems and leaves glabrous, leaf margin lobed as well as toothed 1. B. sonderana 

Stems and leaves hairy, leaf margin toothed but not, or scarcely, lobed 2. B. hirtella 

Placentae entire: 

Leaves symmetrical or nearly so, suborbicular, all, or nearly all, radical 3. B. geranioides 

Leaves mostly asymmetrical, cauline: 

Flowers orange (but often drying pink), male tepals generally 4 4. B. sutherlandii 

Flowers white or pink, male tepals usually 2: 

Primary leaves up to c. 13 x 7 cm, seldom less than 7X3 cm, ratio of length of vein adjacent to main vein in 

broader half of leaf to length of vein running to leaf apex (main vein) 1: (1,1-) 1,3 (-1,5) .5. B. homonyma 
Primary leaves up to 8 x 3,5 cm, more usually c. 5 X 2 cm or less; vein ratio 1: (1,3-) 1,7 (-2,5); 

if in the range 1: 1, 3-1,4, then leaves smaller than the lower limits for B. homonyma 6. B. dregei 

1. Begonia sonderana Irmscher in Bot. 
Jahrb. 81: 156 (1961) (as B. sonderiana). Type: 
Transvaal, Barberton Distr., summit of 
Saddleback Mt ,,Galpin 821 (PRE, holo.; BOL; 
GRA; K; NBG). 

B. caffra sensu Burtt Davy, FI. Transv. 1: 232 (1926). B. 
dregei sensu Burtt Davy, l.c. (1926). B. sonderana var. 
transgrediens Irmscher in Bot. Jahrb. 81: 160 (1961). Type: 
Transvaal, Sabie, Van der Merwe 311 (PRE, holo.).B. sp., 
Letty, Wild Flow. Transv. 204, pi. 101, 2 (1962). 

A glabrous, herbaceous perennial herb, 
often tuberous or the stem decumbent at the 
base, the nodes swollen and rooting. Stems up 
to 1 m tall, stout, fleshy, often pinkish or 
reddish, simple or branched, leafy throughout. 
Stipules lanceolate, membranous, brown. 

Leaves obliquely ovate-acuminate, up to 18 x 
14 cm, but often only about half that size, base 
cordate, margin 5-7-lobed, lobes more or less 
triangular, coarsely and irregularly toothed or 
occasionally crenate, light green and waxy 
above, paler below, veins often reddish, 
petioles up to 15 cm long, often reddish. 
Inflorescences in the upper leaf axils. Flowers 
up to 3 cm across, usually pink, sometimes 
white or white flushed pink, tepals of S flowers 
usually 4, occasionally 2 or 3, of 9 flowers 
usually 5. Fruits 3-winged, more or less 
rounded-oblong or cuneiform in outline, 2X 
2-3 cm across the wings, placentae split, ap- 
pearing bifurcate in transverse section. Figs. 44: 
3; 45: 3. 



Fig. 44. — 1, Begonia geranioides, whole plant, x 4 /s; la, female flower, x 4 /s; lb, ovary, style and stigma, x 2; 1c, ovary 
cut transversely, x 2; Id, stigma, x 5; le, anther, front and side view, x 6; If, seed, x 18 ( Stewart 1640). 2, B. 
sutherlandii, transverse section of ovary with entire placentae (fiilliard & Burn 5647). 3, B. sonderana, transverse 
section of ovary with split placentae (Hilliard & Bunt 5962). 



Ranges from the eastern highlands of Rhodesia and 
Mozambique to the mountains of the eastern and north- 
eastern Transvaal and Swaziland. Favours broken cliff faces 
and rock falls in forest or on forest margins, or shady rock 
outcrops on steep grass slopes between about 1 000 and 
1 850 m above sea level. Flowers mainly from January to 

Transvaal. — 2230 (Messina): Sibasa, Junod sub TRV 
21218. 2330 (Letaba): Woodbush, Codd 9419. 2430 (Pil- 
grims Rest): The Downs, Junod 4194; Mariepskop, Hill- 
iard & Burn 5962; Graskop, Galpin 14443. 2530 (Lyden- 
burg): Sabie, Taylor 1905. 2531 (Komatipoort): 16 km W. 
of Havelock Mine, Codd 6423. 

Swaziland. — 2531 (Komatipoort): near Piggs Peak, 
Compton 32244. 2631 (Mbabane): Sheba’s Veil, Compton 
27278; Hlatikulu, Stewart 111. 

This species was for long confused with B. dregei and 
B. homonyma, but is easily distinguished by its split, not 
entire, placentae. 

2. Begonia hirtella Link, Enum. Hort. 
Berol. 2: 396 (1822). A. DC. in Mart., FI. Bras. 
4: 344 (1861). Type: cult. Berlin, material from 
Brazil (not seen). 

B. villosa Lindl. in Bot. Reg. 15: 1. 1252 (1829). Type: 
cult. hort. R.H.S., seed from Brazil. 

A robust, branched herb. Stems several 
from the base and up to 2 cm diam. there, 
branched, up to 75 cm tall, green, fleshy, 
thickly clothed in long, coarse, white hairs, 
glabrescent. Stipules ovate-lanceolate, about 1 
x 0,5 cm, membranous, whitish, apex acumi- 
nate, margin fimbriate-ciliate. Leaves obliquely 
ovate, up to 10 x 8 cm, apex acute, base 
truncate or nearly so, margins irregularly 
crenate-serrate, not or scarcely lobed, green 
above, thinly villous, paler below, hairs nearly 
confined to the veins, petioles up to 9 cm long, 
fleshy, villous. Inflorescences in the upper leaf 
axils, peduncles up to about 5 cm long, usually 
shorter than the petiole. Flowers up to c. 8 mm 
across, pure white, tepals of 6 flowers 4, of 9 
flowers 5. Fruits 3-winged, rounded-oblong to 
cuneiform in outline, c. 2 cm across the 
broadest part of the wings, placentae split, 
appearing bifurcate in transverse section. Fig. 
45: 2. 

A native of Brazil, long cultivated in Europe, B. 
hirtella appears to have established itself in certain Zululand 
forests, having twice been collected without indication that 
it was not growing naturally. 

Natal. — 2732 (Ubombo): Gwalaweni Forest, Vah- 
rmeijer & Drijfhout 2000. 283 1 (Nkandla): Eshowe, Lawn 

It is an annual, though it will persist for several seasons 
in cultivation, and seeds itself freely. Male and female 

flowers open at the same time, and it is clearly self-fertile, 
so that there is massive seed production. 

Only B. sonderana among native species has split 
placentae like B. hirtella, but is easily distinguished by its 
much larger and generally pink flowers and glabrous, 
distinctly lobed leaves. 

3. Begonia geranioides Hook.f in Bot. 
Mag. 92: t. 5583 (1866); Irmscher in Bot. Jahrb. 
81: 124 (1961). Type: “imported from Port 
Natal by Messrs. Backhouse of York”(K). 

A tuberous, often stemless, perennial herb. 
Leaves several from the crown, petioles up to 
20 cm long, blade suborbicular, up to 8 x 11 
cm, base deeply cordate, apex very obtuse, 
margin often shallowly and irregularly lobed, 
obscurely crenate, both surfaces and generally 
the petiole thinly clad with long white hairs, 
light green, juveniles sometimes white-spotted, 
veins often pinkish or reddish. Flowering stems 
up to 30 cm high, often much shorter, usually 
leafless, occasionally with a few leaves similar 
to but smaller than the radical ones, glabrous or 
sparsely hairy, often pinkish or reddish, bracts 
in pairs subtending each di- or trichotomy, 
ovate, margins sometimes ciliate-fimbriate. 
Flowers 2-3 cm across, pure white, tepals of 6 
flowers generally 4, of 9 generally 5. Fruits 
3-winged, cuneiform in outline, about 2 x 2 cm 
across the broadest part of the wings, placentae 
undivided. Fig. 44: 1; 45: 4. 

Confined to forests between about 600 and 1 375 m 
above sea level from Karkloof in south-central Natal to the 
Zuurberg and Ngeli slopes on the Cape-Natal border near 
Weza. Grows on damp earth banks and rock faces, often 
forming large colonies. Flowers mainly from December to 

Natal. — 2929 (Underberg): Polela, Hlabeni Forest, 
Fernando sub NH 9616. 2930 (Pietermaritzburg): Karkloof, 
Moll 3541; Town Bush Valley, Canham 12; Inanda, 
Groenberg, Johnson 1323; near Richmond, Barker 5230; 
near Ixopo, Crewe 71; Mid Illovo, farm Ismont, Strey 
8354. 3030 (Port Shepstone): near Dumisa, Hilliard & 
Burn 3400. 3130 (Port Edward): Beacon Hill, Strey 6072. 

Cape. — 3029 (Kokstad): Zuurberg, Tyson 1716. 

Easily recognized by its tuft of suborbicular hairy 
leaves. The plant can become dwarfed under dry conditions. 

4. Begonia sutherlandii Hook.f. in Bot. 
Mag. 94: t.5689 (1868); Burtt Davy, FI. Transv. 
1: 232 (1926); Pole Evans in Flow. PI. Afr. 8: 
t.283 (1928); Irmscher in Bot. Jahrb. 81: 162, 
t.8 fig. 1 (1961); Letty, Wild Flow. Transv. 204, 
pi. 101, 3 (1962); Batten & Bokelmann, Wild 
Flow. E. Cape 106, pi. 87, 1 (1966); Wilczek in 
F.C.B. Begoniaceae: 50 (1969). Type: Natal, 
without precise locality, Dec 1861, Sutherland 



Fig. 45. — 1, Begonia sutherlandii, showing leaf variation; la, Nkandla Forest, Zululand (Hilliard & Burn 5647); lb, 
Noodsberg, Natal (Strey 6239); lc, Inanda, Natal ( Wood 1081, type of B. dissecta ); Id, Mapumulo, Natal (Wo// 2945). 
2, B. hirtella, leaf (Vahrmeijer & Drijfhout 2000). 3, B. sonderana (Hilliard & Burn 5962). 4, B. geranioides (IF ood 
749). All X Vs. 



B. sutherlandii Hook.f. var. latior Irmscher in Bot. Jahrb. 
81: 165, t.9 figs. 1 & 2 (1961), forma latior Irmscher, l.c. 
Type: Zambia, above Mukoma, Inono Falls, 900 m, 
Richards 3699 (B, holo., not seen: K). B. dissecia Irmscher, 
1. c. 81: 178, 1. 1 1, fig. 2 (1961). Type: Natal, Inanda, Wood 
1081 (K, holo.; NH; NBG). B. buttonii Irmscher, l.c. 81: 
178 (1961). Type: Natal, Durban, (Port Natal) Button 8 (K). 
B. suffruticosa Meisn. var. gueinziana A. DC. in DC., Prodr. 
15: 385 (1864). Type: Natal, Durban (Port Natal), Guein- 
zius 248 (W, not seen). B. gueinziana (A. DC.) Irmscher, 
l.c. 81: 177 (1961). 

An herbaceous perennial with one or sev- 
eral stems from a tuber. Stems up to 1 m tall, 
simple or branched, slender, fleshy, leafy 
throughout, they and the leaves glabrous to 
thinly hairy, green or partially or wholly tinged 
red as are the stipules, petioles, veins and 
sometimes the leaf margins as well. Stipules 
lanceolate to ovate, margins often fimbriate- 
ciliate. Leaves very oblique, elliptic-lanceolate 
to ovate, up to 25 x 15 cm, but often only 
two-thirds or less that size, base cordate to 
truncate, apex acuminate to long-acuminate, 
margin acutely and irregularly serrate, fre- 
quently lobed as well, or occasionally more 
deeply cut, thin-textured, light bright green 
above with a satiny sheen, paler below, petioles 
slender, up to c. 15 cm long. Inflorescences 
axillary and terminal, many-flowered. Flowers 
up to 3,5 cm across, very showy, varying from 
pale to deep orange or brick red (often drying 
pink), tepals of S flowers 4, of $ flowers usually 
5. Fruits 3-winged, cuneiform in outline, up to 
2 x 3 cm across the broadest part of the wings, 
placentae entire. Fig. 44; 2; 45: 1. 

One of the most widespread of the African Begonias, 
ranging from Katanga, Tanzania, Zambia and the eastern 
highlands of Rhodesia through the mountains of the eastern 
Transvaal and Swaziland to Natal, where it is common in 
forest patches up to about 1 900 m above sea level, and into 
the eastern Cape (Transkei) as far south as Kei Mouth. 
Grows in forest, on the steeply sloping forest floor or on 
earth banks, rock faces or mossy boulders, sometimes 
epiphytic, often forming large colonies. Flowers mainly in 
December and January. 

Transvaal.— 2430 (Pilgrims Rest): The Downs, Junod 
4183; Mariepskop, Hilliard & Burtt 5969. 2530 (Lyden- 
burg): Dullstroom, Galpin 13731; Lunsklip Falls, Codd 
10013. 2531 (Komatipoort): Barberton, Galpin 727. 2630 
(Carolina): Ermelo, Collins sub TRV 6531 . 

Swaziland. — 2531 (Komatipoort): near Havelock, 

Compton 29845. 

Natal. — 2730 (Vryheid): Nyati Mt., Hilliard & Burtt 
5897. 2829 (Harrismith): Oliviershoek Pass, Hilliard 4910; 
Tintwa Mts, Doidge sub PRE 31569. 2831 (Nkandla): 
Nkandla Forest, Hilliard & Burtt 5647; Eshowe district, 
Gerstner 3204. 2929 (Underberg): Bulwer, Marwaqa Mt., 
Marais 1448; Cathkin Park, Galpin 11764. 2930 (Pieter- 
maritzburg): Umgeni Poort, Moll 1403; Little Noodsberg, 

Strey6250; Table Mt., McClean 152. 2931 (Stanger): near 
Mapumulo, Moll 2945. 3029 (Kokstad): Ngeli Mt., Hill- 
iard & Burtt 5753. 3030 (Port Shepstone): farm Ellesmere, 
Strey 7054; Paddock, Strey 6193. 3130 (Port Edward): 
Beacon Hill, Strey 6070. 

Cape. — 3028 (Matatiele): near Mt. Frere, Story 947. 
3029 (Kokstad): Tabankulu Mt., Hilliard & Burtt 6526; 
Fort Donald, Tyson 1655. 3128 (Umtata): Baziya Mt., Baur 
391 . 3129 (Port St. Johns): Port St. Johns, Flanagan 2516. 
3228 (Butterworth): Kei Mouth, Flanagan 419. 

There is considerable variation in stature, leaf size and 
leaf cutting, in indumentum, and in the presence or absence 
of red tints in the vegetative parts of this species, the most 
handsome and, by virtue of its beautiful orange flowers, the 
most easily recognized of the South African Begonias. 
Favouring as it does banks, cliffs and boulders, it is very 
susceptible to the effects of drought, and there is little doubt 
that environmental factors profoundly influence stature and 
leaf size. There seems to be some correlation too between 
light intensity and the development of the red coloration in 
stems, petioles, veins and so on. The degree of lobing and 
cutting of the leaf margin is more probably under genetical 
control. The margin is always sharply serrate and may be 
shallowly to deeply lobed, even deeply dissected, but the 
variation is continuous. Nearly all specimens from the 
Transvaal and Swaziland are noticeably hairy with both leaf 
surfaces, and often the petiole as well, clothed in long (up to 
3 mm) hairs, while in Natal, the Transkei and eastern Cape, 
leaves are generally glabrous or have a sparse covering of 
very short hairs. However, around Sabie and Barberton in 
the Transvaal, plants may be hairy or nearly glabrous, while 
specimens with long hairs on the leaf surfaces but scarcely 
extending onto the petiole occur in widely separated areas in 
Natal and the Cape, for example, in Ndwedwe district 
(Strey 6131; 6064; 6250), Umzinto district (Rudatis 811; 
Strey 7054) and at Port St. Johns in the Transkei (Flanagan 
2516 and Flanagan sub herb. Bolus 8717). 

5. Begonia homonyma Steud., Nom. 
Bot. ed. 2: 194 (Aug. 1840); B.L. Burtt in Notes 
Roy. Bot. Gard. Edinb. 32: 274 (1973). Type: 
cult. Edinb. ex Berlin. 

B. sinuata E. Mey. ex Otto & Dietr. in Allg. Gartenztg. 4: 
357 (5 Nov. 1836), nom. illegit. , non Wall, ex Meisn. (Aug. 
1836). Type: cult. Berlin (not seen). B. sinuata Grah. in 
Edinb. New Phil. Joum. 24: 192 (1838); in Bot. Mag. 
t. 373 1 (1839), nom. illegit., non Meisn. (1836). Type as for 
B. homonyma. B. caffra Meisn. in Linnaea 14: 501 (1841); 
Sond. in F.C. 2: 481 (1862); Irmscher in Bot. Jahrb. 81: 136 
(1961). Type as for B. sinuata Otto & Dietr. B. dregei Otto 
& Dietr. var. sinuata A. DC. and var. caffra A. DC. in DC., 
Prodr. 15: 384 (1864). B. favargeri Rechinger in Ann 
Naturh. Hofmus. Wien 20: 33 tab. 1 (1905); Thonner, FI. 
PI. Afr., 1. 107 (1915). Type: Natal, at the Umkomaas River, 
Krook (PI. Penther. no. 2976, W, not seen). B. rudatisii 
Irmscher in Bot. Jahrb. 81: 129 (1961). Type: Natal, 
Umzinto distr., Dumisa, Friedenau, Rudatis 602 (B, not 
seen; E; K). 

Augustia caffra (Meisn.) Klotzsch, Beg. Gatt. Art.: 81 

A glabrous, herbaceous perennial herb. 
Stems erect, 1 or several from a large tuber, up 
to 1 m tall, simple or branched, stout, fleshy, 
sometimes tinged red, leafy throughout. 



Stipules oblong-lanceolate, membranous, 
brown. Leaves obliquely ovate, up to c. 13 X 7 
cm, base shallowly cordate, apex broadly acute, 
margins entire, sinuate or broadly and shallowly 
lobed, the lobes rarely subserrate, thin-textured, 
crisp, light green and waxy above, paler below, 
the young ones often white-spotted, veins (in 
dried material) strongly raised below particu- 
larly near the basal sinus, often reddish, petioles 
up to 13 cm long, but often of the order 3-5 cm, 
decreasing in length upwards, sometimes red- 
dish. Inflorescences in the upper leaf axils. 
Flowers up to 3 cm across, white, pink or white 
tinged pink, tepals of 6 flowers usually 2, of 9 
flowers usually 5. Fruits 3-winged, about 2X2 
cm across the expanded wings, placentae entire. 
Fig. 46: 1. 

Recorded from the more coastal districts of Natal and 
the Transkei, up to about 900 m above sea level, from the 
southern end of the Lebombo Mts. to the Great Kei, on rock 
falls and cliffs in forest. Flowers from December to March. 

Natal. — 2732 (Ubombo): Gwalaweni Forest, Vah- 
rmeijer & Drijfhout 2002 . 2930 (Pietermaritzburg): Bothas 
Hill, McClean 116. 2931 (Stanger): Umvoti Valley, near 
Glenmill, Hilliard 5084. 3029 (Kokstad): near Gundrift, 
Acocks 12237. 3130 (Port Edward): Umtamvuna Bridge, 
Strey 5982. 

Cape. — 3029 (Kokstad): Clydesdale, Tyson 2689. 3129 
(Port St. Johns): near Lusikisiki, Strey 8930. 3228 (Butter- 
worth): Collywobbles, Van Breda 878. Grid uncertain: 
Gcalekaland, N. of Great Kei River, Dinnie 665. 

B. homonyma together with B. dregei and the forms 
placed in synonymy thereunder represents a highly variable 
and complex group. B. homonyma can, however, be distin- 
guished by its large obliquely ovate leaves with entire or 
shallowly lobed margins and veins strongly raised below 
particularly near the basal sinus. There is an almost clearcut 
discontinuity in the absolute size of the mature primary 
leaves of B. homonyma and B. dregei: in B. homonyma it 
scarcely falls below 7 x 3 cm and is more usually about 1 1 
x 6 cm; in B. dregei the largest leaf measured was 8 X 3,5 
cm, but more usually they are about 5 x 2 cm or less. 

A more reliable measurement (one that must largely 
circumvent the effects of environmental factors and is an 
indicator of leaf shape rather than size) is the ratio obtained 
by dividing leaf breadth into leaf length (length of the vein 
running out to the leaf tip in this case = length, and the 
length of the adjacent vein in the broader half of the leaf = 
breadth). In B. homonyma this is 1: (1,1—) 1,3 (—1,5); in B. 
dregei 1: (1,3-) 1,7 (-2,5). The lower ratios in B. dregei (1: 
1,3— 1,4) are from small, deeply serrate or partly dissected 
leaves that cannot be confused with those of B. homonyma. 

6. Begonia dregei Otto & Dietr. in Allg. 
Gartenztg. 4: 357 (1836); Meisn. in Linnaea 14: 
502 (1840); Sond. in F.C. 2: 481 (1862); A.DC. 
in DC., Prodr. 15: 384 (1864), excl. var. sinuata 
and var. caffra; Irmscher in Bot. Jahrb. 81: 130 
(1861). Type: Cape, between the Umtata and 
Umgaziana rivers, Drege (B, not seen). 

B. parvifolia sensu Graft, in Edinb. New Phil. Joum. 
xxiv: 91 (1838); in Bot. Mag. t.3720 (1839). B. sujfruticosa 
Meisn. in Linnaea 14: 502 (1840); Sond. in F.C. 2: 481 
(1862); A.DC. in DC., Prodr. 15: 385 (1864) excl. var. 
gueinziana A.DC.; Irmscher in Bot. Jahrb. 81: 130 (1961), 
inch forma holusii Irmscher and forma worsdellii Irmscher. 
Type: Cape, between the Umgaziana and Umzimvubu 
rivers, Drege 8032 (B, not seen). B. natalensis Hook, in 
Bot. Mag. t.4841 (1855); Sond. in F.C. 2: 481 (1862); 

A. DC. in DC., Prodr. 15: 385 (1865); Irmscher in Bot. 
Jahrb. 81: 138 (1961). Type: the plate in Bot. Mag., from 
specimen collected by Capt. Garden in Natal, not preserved. 

B. richardsiana T. Moore in Gard. Chron. 1065 fig. 243 
(1871); Regel in Gartenflora 21: 293 tab. 739 fig. b (1872); 
Houllet in Rev. Hort. 44: 333 fig. 35 (1872), sphalm. 
richardsoniana; R. A. Dyer in Flow. PI. Afr. t.673 (1937). 
Type: spec. cult. R.H.S. (K). B. partita Irmscher in Bot. 
Jahrb. 81: 143, tab. 10, fig. 2 (1961). Type: Natal, Umzinto 
district, farms Campbellton and Glenrosa, 550 m, Rudatis 
1876 (B, holo, not seen; NBG). 

Augustia dregei (Otto & Dietr.) Klotzsch, Beg. Gatt. Art. 
80 (1855). A. natalensis (Hook.) Klotzsch, l.c. 81 (1855). 
A. sujfruticosa (Meisn.) Klotzsch, l.c. 81 (1855). 

Glabrous (or occasionally a few fleshy setae 
on the upper leaf surface) perennial herbs. 
Stems up to 30 cm tall, one or several from a 
tuber, simple or branched, erect or sometimes 
decumbent at the base and there rooting, thick, 
fleshy, sometimes reddish, leafy throughout. 
Stipules oblong-lanceolate, membranous, 
brown. Leaves obliquely-ovate to obliquely 
ovate-lanceolate, up to 8 x 3,5 cm but more 
usually about 5x2 cm, base shallowly cordate 
to subtruncate, apex narrowly triangular-acute 
to long-attenuate and acuminate, margins rarely 
subentire to sinuate, more usually more or less 
regularly lobed or with 1 or 2 spreading lobes in 
the lower part of the broader half of the leaf, the 
narrower half lobed or not, the lobes in turn 
coarsely serrate, sometimes the leaf more 
deeply or even relatively finely dissected, 
somewhat fleshy, light green above, paler be- 
low, the young leaves often, the mature occa- 
sionally, white-spotted, veins green or reddish, 
not, or very rarely, raised below, petioles up to 
9 cm long, often 4—5 cm, always decreasing in 
length upwards, green or reddish. Inflores- 
cences in the upper leaf axils. Flowers up to 3 
cm across, white, pink, or white tinged pink, 
tepals of 6 flowers usually 2, of 9 flowers 
usually 5. Fruit 3-winged, 1-2 x 1-2 cm 
across the expanded wings, placentae entire. 
Fig. 46: 2. 

B. dregei ranges from Qudeni, Ngotshe and Ngoye 
forests just north of the Tugela in Zululand, through coastal 
Natal as far south as East London in the eastern Cape, 
usually below 600 m altitude. It therefore occupies virtually 
the same area as B. homonyma. Like that species, and other 
South African Begonias, it grows on rocky cliffs, steep 



Fig. 46. — To show leaf variation in 1, Begonia homonyma and 2, B. dregei; la and lb both from the Umvoti river. Natal 
( Hilliard 5084). 2a, Port St. Johns, Transkei (Admiraal 2286); 2b, Port St. Johns ( Strey 6655); 2c and 2d, both on one 
specimen from the Bashee River mouth, Transkei ( Gordon-Gray 1564); 2e, Mqanduli, Transkei ( Theron 1504); 2f, Kei 
Mouth, Transkei ( Flanagan 419); 2g, Cobe, Transkei (Pegler 813); 2h, Port St. Johns (Sim 20231); 2i, Kentani, 
Transkei (Pegler s.n.); 2j, hort. Kew Nov. 1860; 2k, Inanda, Natal (Wood s.n.); 21, Eshowe, Zululand (Lawn 1447); 
2m, Lupatana, Transkei (Strey 10223); 2n, Inanda (Wood 1197); 2o, Ngotshe Forest; Eshowe (Hilliard &Bum 5657); 
2p, Ngoye Forest, Zululand (Wood 10349); 2q, Ngoye Forest (Hilliard & Burn 5640); 2r and s, both on a single 
specimen from Coffee Bay, Transkei (Wells 3539); 2t, Qora Mouth, Transkei (Wells 3595); 2u, Port St. Johns (Bolus 
8914); 2v, Port St. Johns (Mogg s.n ); 2w, Manubi Forest, Transkei (Worsdell s.n.). 2a is the form described as B. 
dregei, 2j as B. natalensis, 2p as B. partita, 2u and 2v as B. suffruticosa. All x 3 /s. 



earth banks and among rock falls in forest, and flowers 
mainly from December to March. Although the material 
may be classified into several groups, it is preferred not to 
adopt a formal infraspecific nomenclature for them at this 

GROUP 1 ( dregei sens, strict.) 

Natal. — 2831 (Nkandla): near Eshowe, Lawn 517. 

Cape. — 3129 (Port St. Johns): Port St. Johns, Admiraal 
2286; Mqanduli district, banks of Umnenga River, Theron 
1504 . 3228 (Butterworth): Kei Mouth, Flanagan 419. 

GROUP 2 ( dregei — * natalensis ) 

Natal. — 3030 (Port Shepstone): Paddock, Strey 5886; 
St. Michaels-on-Sea, Ross 1885. 

Cape. — 3129 (Port St. Johns): Port St. Johns, Sim20231. 
3228 (Butterworth): Kentani, Pegler 813; Bashee River 
Mouth, The Haven, Gordon-Gray 2006. 3327 (Peddie): 
East London, Batten s.n. 

GROUP 3 (natalensis sens, strict.) 

Natal. — 2830 (Dundee): Qudeni, Gerstner 3233. 2831 
(Nkandla): near Eshowe, Hilliard & Burn 5657 (leaves 
here occasionally bilobed and thus approach B. partita; a 
similar specimen, Lawn 1447 from near Eshowe, was 
designated partita X natalensis by Irmscher). 2930 
(Pietermaritzburg): Inanda, Wood 1197. 3030 (Port 

Shepstone): farm Excelsior, Hilliard & Burn 3390. 

Cape. — 3129 (Port St. Johns): near Lusikisiki, Strey 
10198; Lupatana, Strey 10223. 3227 (Stutterheim): Horse- 
shoe, Galpin 8212. 

GROUP 4 (partita sens, strict.) 

Natal.— 2831 (Nkandla): Ngoye, Wood 10349; Hill- 
iard & Burn 5640. 3030 (Port Shepstone): Port Shepstone, 
Dimock Brown 489. 

GROUP 5 ( suffruticosa sens, strict.) 

Cape. — 3129 (Port St. Johns): Port St. Johns, Bolus 
8914; Flanagan 2510; Galpin 3465; Howlett 21; Mogg 
20157; Moss 4620; Schelpe 285; Schonland 4185. 

The extensive synonymy of B. dregei reflects the high 
degree of variability in leaf lobing and cutting and, to a 
lesser extent, in leaf shape. The specimens cited have been 
grouped to indicate this variation and the associated names. 
The variation is, however, continuous and is complicated by 
the fact that juvenile and secondary leaves are often entire 
or less deeply cut than mature primary leaves. For instance, 
in Gordon-Gray 2006 (cited in Group 2 above) some of the 
leaves, particularly the lower primaries, fit the criteria for B. 
dregei, while upper and secondary leaves fit those of B. 

Graham illustrated as B. parvifolia (Bot. Mag. t.3720) 
a plant received from the Berlin garden in 1836, the year in 
which B. dregei was described. The plants were undoub- 
tedly the same and probably of the same stock. The Bot. 
Mag. plate may therefore be taken to depict “typical” B. 
dregei, having small obliquely ovate leaves, apex broadly 
acute, margin more or less regularly lobed, the lobes 
toothed. This leaf form grades imperceptibly into that of 
typical B. natalensis: leaf more elliptic-ovate in outline, 
with two broad lobes in the basal half and a long-attenuated 
upper half, margin typically serrate (see Bot. Mag. t.4841) 
but grading to sinuate or entire. Irmscher designated some 
intermediate plants as hybrids between B. dregei and B. 
natalensis, but they seem to be merely part of the continu- 
ous variation pattern (see fig. 46). 

The nam e. partita is attached to a specimen with deeply 
bilobed leaves, these two main lobes further lobed, margins 
entire or serrate. B. suffruticosa forma worsdellii has the 
deep bilobing of B. partita but in the marginal toothing 
begins to approach B. suffruticosa. B. suffruticosa is 
represented in herbaria by eight collections, all from Port St. 
Johns. The type locality “between the Umgaziana and 
Umzimvubu rivers” is also Port St. Johns. There seems 
little doubt that B. suffruticosa is merely a local variant with 
deeply dissected leaves, linked through B. suffruticosa 
forma worsdellii and other variants to B. partita and so to 
B. natalensis and B dregei. 

B. dregei is very closely related to B. homonyma, and 
the whole complex demands a thorough field and laboratory 
study. For that reason, no infraspecific categories are 
recognized here. 

Both B. dregei and B. homonyma have been much 
confused with B. sonderana, a more northern species easily 
distinguished by its split placentae. 


by A. A. Obermeyer 

Perennial, herbaceous, suffruticose or arborescent; predominantly succulents, rarely epiphy- 
tic. Stems woody or mostly succulent and variously modified, cylindrical, flattened or globose, 
bearing axillary, rounded, usually raised, woolly areoles which bring forth leaves, flowers, 
spine-clusters and often barbed bristles. Leaves present, usually early deciduous, often reduced or 
0. Flowers solitary or rarely clustered, sessile (except Pereskia), bisexual, mostly regular. 
Hypanthium (receptacle-tube) long or short, naked or with leaf-like bracts and areoles containing 
reduced spines, wool and bristles and giving rise to a series of imbricate, sepaloid green segments 



which gradually change into a series of often very showy petaloid segments. Stamens numerous, 
inserted on the receptacle, free or adnate to the petaloid segments, anthers bilocular, opening 
lengthwise. Ovary inferior, 1-locular with 3 or more, many-ovuled, parietal placentas; style 
simple; stigmas 3-many. Fruit usually a berry, often edible, rarely a capsule; seeds few to 

Genera 84, species approximately 2 000; widespread in the Americas with many naturalized and often troublesome 
elsewhere. A great many species representing a number of genera are grown by gardeners and hobbyists but only those 
which have become naturalized are dealt with here; most of these have been proclaimed as noxious weeds. One genus, 
Rhipsalis, appears to be indigenous. 

Plants with succulent stems variously shaped; leaves 0 or few and then usually reduced and 
early deciduous; flowers solitary in each areole (rarely 2 or more in Rhipsalis but then 

Glochidia (barbed bristles) absent; stems terete, succulent, not articulated, supple when 
young; leaves absent; seeds black or brown, not encased in a long aril: 

Terrestrial, large clambering plants without aerial roots; stems c. 2 cm in diam.; flowers 

c. 20 cm long, funnel-shaped 1. Harrisia 

Epiphytic or saxicolous plants with aerial roots; stems 3-6 mm in diam.; 

flowers c. 5 mm long 2. Rhipsalis 

Glochidia present inside areoles; stems firm, terete or flattened, often articulated; leaves 
rarely present, usually small and early deciduous; seeds encased in a bony, white 

aril 3. Opuntia 

Plants with normal stems and leaves; flowers large, several from each areole 4. Pereskia 



Harrisia Britton in Bull. Torr. Club 35: 561 (1908); Britton & Rose, The Cactaceae 2: 147 (1920); 
Hunt in Hutch., Gen. Flow. PI. 2: 445 (1967). Type species: H.(Cereus) gracilis (Mill.) Britton. 

Cereus, group 18, Eriophori, A, Euharrisia, Vaupel in Pflanzenfam. ed. 2, 21: 633 (1925). 

Arborescent or clambering shrubs with long supple 3-12-ribbed or -angled stems, bearing several 
usually stout spines in each areole. Flowers nocturnal, solitary, large, funnel-shaped; outer 
segments linear to lanceolate, greenish or pink, inner white or pink; the cylindric hypanthium as 
long as the limb or longer, bearing pubescent areoles and scales; stamens numerous; ovary and 
young fruits tubercled. Fruit globose to obovoid-globose, with or without spines, scales deciduous; 
seeds numerous, small, dull black, somewhat verrucose. 

About 20 closely related species recorded from tropical America to Argentina. One species recently naturalized in 

Harrisia martinii (Labouret) Britton in 
Addisonia 2: 55 (1917); Britton & Rose, The 
Cactaceae 2: 155, 1. 19, f.3, t. 20, f.2 (1920). 
Type: probably not preserved; origin unknown. 
Recorded from Argentina. 

Cereus martinii Labouret, Ann. Soc. Hort. Haute 
Garonne (1854). 

Plants with tuberous roots and long, fairly 
supple, clambering, much branched stems, 1-2 
cm in diam., 4—5-angled; areoles numerous, 
placed fairly close together, raised and shortly 
pubescent bearing a stout central, patent or 

reflexed spine up to 3 cm long and several very 
short radiating spines. Flower about 20 cm 
long; outer segments linear, greenish; inner 
broader, white or pinkish. Fruit globose, 3,5 
cm in diam., red, with small scales, the with- 
ered perianth long persistent; seeds small, 
black, scattered in a white pulp. Fig. 47. 

Recorded from Natal in the neighbourhood of Pieter- 
maritzburg where it has become troublesome in some areas, 
the birds transporting the seeds. Declared a weed through- 
out the Republic in 1968. 

Natal. — 2930 (Pietermaritzburg): Ashburton, Regional 
Officer sub PRE 32235; Paterson sub PRE 32308. 



Fig. 47. — 1, Harrisia martinii, portion of flowering branch, x 2 /j; a, fruit, x 2 6 (After Britton & Rose). 



5416 2. RHIPSALIS 

Rhipsalis Gaertn., Fruct. Sem. 1: 137, t. 28 (1788) nom. conserv.; Britton & Rose, The Cactaceae 
4: 219 (1923); Vaupel in Pflanzenfam. ed. 2, 21: 617 (1925); Phill., Gen. ed. 2: 522 (1950); Hunt in 
Hutch., Gen. Flow. PI. 2: 427 (1967), in F.T.E.A. Cactaceae: 5 (1968). Type species: R. baccifera 
(J. Mill.) Steam ( =R . cassutha Gaertn.). 

Unarmed succulents, often epiphytic. Stems pendulous, branching freely, terete, angled 
and/or jointed or flattened and leaf-like, areoles small, woolly, bristly, rarely spinous. Leaves 
reduced to minute scales or 0. Flowers usually solitary, lateral, sessile, small. Perianth segments 
5-12, with the outer sepaloid series short and free or nearly so, and the inner petaloid, fused below, 
mostly white. Stamens few to numerous, inserted at the base of the perianth or attached to the base 
of the petals. Ovary usually inferior with 3-several parietal placentas. Berry globular, translucent, 
smooth or with some scattered, reduced areoles containing bristles or spines, juice viscid; seeds 
oblong, irregularly angled, brown to black. 

About 50 species in South America, Mexico and West Indies. A few of these naturalized elsewhere; one species, 
probably R. cereusculus Haw., is often cultivated in South Africa. The following species, doubtfully native, found wild in 
moist warm regions of Africa, Madagascar, Mascarenes and Sri Lanka. 

Rhipsalis baccifera (J. Mill.) Steam in 
adnot. Cactus J. 7: 107 (1939); Hunt in 
F.T.E.A., Cactaceae: 5, fig. 2 (1968); Troupin, 
Syll. FI. Rwanda, Spermatophytes: VI, 47 with 
figure (1971). Type: description and plate, 
Class IX, Ord. 1 (1771) in J. Mill., Illustr. Sex. 
Syst. Linn. (1771-77). 

Cassyta baccifera J. Mill., Illustr. Sex. Syst. Linn. Class 
IX, Ord. 1 (1771-77). Type as above. 

R. cassutha Gaertn., Fruct. Sem. 1: 137, t.28 (1788); 
Hook, in Bot. Mag. 58: t.3080 (1831), as R. cassytha, the 
spelling subsequently adopted by most authors; Oliv. in 
F.T.A. 2: 581 (1871); Britton & Rose, The Cactaceae4: 225 
(1923); Vaupel in Pflanzenfam. ed. 2, 21: 619 (1925); Burtt 
Davy, FI. Transv. 1: 237 (1926); Bally in J. E. Afr. Nat. 
Hist. 16: 44 (1942); Roland-Gosselin in Desert Plant Life 
1947: 121 (1947). Type: ex hort. Kew, probably not 
preserved; origin unknown. 

Epiphytic or saxicolous, small succulents 
with much branched, pendulous, succulent, ter- 
ete, soft, glaucous-green stems about 3-6 mm 
in diam., bearing adventitious aerial roots co- 
vered with a thin velamen; branches clustered, 
articulated; areoles scattered, woolly, bearing 
few to many soft bristles up to 4 mm long which 
usually disappear with age but the juvenile form 
may persist for many years. Flowers 1-2 per 
areole, sessile, small, green, self-fertile; hypan- 
thium bulbous, bearing a few reduced areoles 
with 1-2 soft short bristles; sepaloid segments 
few, minute, triangular; petaloid segments 4—6, 
oblong, irregular, c. 3 mm long, somewhat 
fleshy, greenish- white. Stamens 5-10, placed 
on receptacle, short, with short opposing 

locules, apiculate. Ovary embedded in bulbous 
hypanthium; style short. Berry globose or 
oblong-globose, 5-10 mm in diam., translucent, 
greenish-white, white, pink or red, the petaloid 
segments semi-persistent, sap viscid; seeds ob- 
long, irregularly and obtusely angled, 1 mm 
long, black or dark brown, shiny, reticulate. 
Fig. 48. 

Widespread in South and Central America; in moist 
warm regions of tropical west Africa to south eastern 
Africa, Madagascar, Mascarenes and Sri Lanka. Plentiful in 
Zululand, southern Natal and north eastern Cape, in coastal 
evergreen forests on trees or on rocks. Distributed by birds 
which are attracted to the berries; these contain a viscid 
juice which aids dispersal. 

Swaziland. — 2631 (Mbabane): Isateki Beacon south of 
Stegi, Compton 27900; 27316. 

Natal. — 2731 (Louwsburg): Lebombo Mts., Border 
Cave, 1 Veils 4456; Pongola Poort, Repton 5982; Strey 
4645; Josini Dam, Codd 10313. 2831 (Nkandla): Umfolozi 
Game Reserve, Bourquin 449a; Eshowe-Entumeni Road, 
Gerstner sub NH 22449; Lawn 435; Ngoye Forest, Wells 
& Edwards 51. 2832 (Mtubatuba): Hluhluwe Game Re- 
serve, Ward 2057; False Bay, Ward 1635. 2930 (Pieter- 
maritzburg): Inchanga Valley, Van der Merwe 2843. 2931 
(Stanger): Thrings Post, Moll 2305. 3030 (Port Shepstone): 
Paddock, Me Clean 324; Strey 6268; Kenterton, along Ifafa 
River Valley, Bridgman sub PRE 32181; Mgongongo, 
Strey 8097. 3130 (Port Edward): Beacon Hill East, Strey 

Cape. — 3129 (Port St. Johns): Waterfall Bluff, 
Lusikisiki, Van der Merwe 2515; Luputana, Strey 10235; 
Port St. Johns, Flanagan 2598; West Gate, Galpin 3197. 
3228 (Butterworth): 1 1 km from coast on Kei Mouth road in 
Kei River Valley towards Komga, Dyer 4506. 



Fig. 48. — 1, Rhipsalis baccifera, habit, x 1 ( Vahrmeijer & Hardy 1692); a, flower, x 6; b, berry, x 6; c, seed, x 40; d, 
seedling, showing' juvenile pubescence. 




Opuntia Mill., Gard. Diet. Abr. ed. 4 (1754); Britton & Rose, Cactaceae 1: 42 (1919); Vaupel in 
Pflanzenfam. ed. 2,21: 647 (1925); Hunt in Hutch., Gen. Flow. PI. 2: 435 (1967); F.T.E.A. 
Cactaceae: 1 (1968). Lectotype species: O. compressa (Salisb.) Macbr. (Cactus opuntia L., Sp. PI.: 
468, 1753, partly). For synonyms see Hunt in Hutch., Gen. Flow. PI. 2: 435 (1967). 

Arborescent, suffruticose or stoloniferous, gregarious, perennial succulents usually spreading 
through breakage of brittle joints (i.e. jointed succulent green branch-segments). Roots fibrous 
and/or tuberous. Stems in arborescent plants forming woody, cylindrical trunks; the joints 
cylindrical or flattened, linear to rounded, usually contracted below at the node, glabrous or 
pubescent, beset with few to many scattered areoles (metamorphosed axillary buds) which are 
variously pubescent, bear barbed bristles (glochidia) and usually one to many radiating, unequally 
long spines, besides secondary joints, leaves and flowers which are usually placed apically. Leaves 
when present mostly small, terete or subulate, usually early deciduous. Flowers solitary in each 
areole, fairly large with a green hypanthium (modified joint) beset with few to many areoles; 
sepaloid segments usually small, green, bract-like, grading into many yellow, orange or red, often 
showy, erect or spreading petaloid segments; stamens numerous; ovary embedded in the swollen 
hypanthium; style terete, stout, stigma-lobes 6-8. Fruit a berry, the epicarp fleshy, ovoid to 
globose, often with a concave apex; areoles few to many, the spines absent, few, or reduced to 
bristles; seed discoid or reniform, covered by a hard funicular aril, embedded in a soft juicy 

About 240 variable species in North and South America. A small number have been in cultivation in the Old World for 
centuries and were used for food, forage and protective hedges. In recent times a number of garden escapes have become a 
terrible scourge in many countries where they have invaded large tracts of land. Their eradication is proving very costly. At 
present the most serious Opuntia infection is found in the eastern Cape, where O. aurandaca and O. ficus-indica are 
invading more and more land and presently cannot be controlled. 

A number of spineless hybrids from the United States have been introduced as stock feed in regions often plagued by 

Joints cylindrical, tuberculate: 

Leaves long-persisting, arcuate, 5-12 cm long; spines not sheathed, 1-2 per areole; flowers orange or greenish 

yellow 1.0. exaltata 

Leaves early deciduous or absent, up to 2,5 cm long; spines covered with papery sheaths, 10-20 per areole; flowers 

Spines close together almost hiding the joint, whitish; small compact bushes up to 60 cm tall 2. O. rosea 

Spines more widely spaced, light brown; open branching shrubs, 1-3 m tall 3. O. imbricata 

Joints flattened or semi-terete, not tuberculate: 

Low spreading plants with semi-terete, densely spiny joints, 1,5-2, 5 cm wide 4. O. aurandaca 

Arborescent or shrubby with flat, rounded to narrowly obovate joints more than 6 cm wide: 

Joints orbicular, about as long as broad: 

Joints with close-set short dark thin spines c. 15 mm long, which are mostly deflexed: hypanthium cup-shaped, 

c. 2 cm long 5. O. spinulifera 

Joints with some scattered atrophied areoles, the upper bearing 1-2 hard, flattened, slightly curved spines 3-5 

cm long; hypanthium cylindrical, c. 4 cm long 6. O. aff. O. lindheimeri 

Joints obovate, attenuate towards the base: 

Areoles large, over 1 cm in diam., bearing 4—7 coarse hard brown spines up to 5 cm long, emerging from a 

bristly cushion; hypanthium and fruit curved l.O. dillenii 

Areoles smaller, about 5 mm in diam., with fewer, thinner greyish spines or unarmed; hypanthium and fruit 

Low spreading bushes 1-2 m tall; fruit obovoid, smooth, purple S O. stricta 

Taller plants, often arborescent, forming trunks; fruit ovoid to obovoid with many scattered bristly areoles, 
red, yellow or purple: 

Spines c. 1 or 2 per areole, 3-7 cm long; joints usually thin, bright green and shiny at least when young, 

the margin receding between the areoles in dried specimens; fruit obovoid, purplish red .9. O. vulgaris 
Spines 3-10 per areole, up to 3 cm long or absent in old plants; joints fairly thick, glaucous with straight 

margins; fruit ovoid to somewhat barrel-shaped, reddish or yellowish 10. O. ficus-indica 



1. Opuntia exaltata A. Berger, Hort. 
Mortol. 410 (1912); Britton & Rose, The Cac- 
taceae 1: 76 (1919); Hunt in F.T.E.A. Cac- 
taceae: 2 (1968). Type: Described from living 
plants in Sir Thomas Hanbury’s garden at La 
Mortola, Italy. 

Much-branched spreading shrubs forming 
thickets, developing a trunk with age. Joints 
arcuate, cylindrical, tuberculate, c. 3 cm in 
diam., glaucous-green. Leaves linear to terete, 
3-6 (-12) cm long, arcuate, fleshy, persistent. 
Areoles with a protruding white woolly pubes- 
cence; with 1 or 2 acicular, straight, strong 
yellow-brown spines up to 5 cm long. Flowers 
with the hypanthium forming a reduced apical 
spineless joint which in turn may proliferate; 
petaloid segments 2-3 cm long, orange or 
greenish yellow. Fruit not seen in South Afri- 
can material. (“Green, pear-shaped, 9 cm long, 
usually sterile; seeds large, irregular, 10 mm 
broad’’ fide Britton & Rose). 

Recorded as a pest from the Orange Free State, Cape 
and Natal. Found both wild and cultivated in Ecuador, Peru, 
Bolivia and probably northern Chile, according to Britton & 

Transvaal.— 2528 (Pretoria): Pretoria, Gericke sub 
PRE 31403. 2627 (Potchefstroom): On farm Leeuwfontein, 
Potchefstroom district, Du Toil sub PRE 31402. 

O.F.S. — 2827 (Senekal): Ficksburg district, Dreyer sub 
PRE 24037. 2926 (Bloemfontein): Bloemfontein district, 
Dreyer sub PRE 24036. 

Natal. — 3030 (Port Shepstone): Umzimkulu, Du Toit 
sub PRE 32673. 

Cape. — 3318 (Cape Town): Koelenhof, on a farm. 
Cars tens sub PRE 32672. 

Specimens at PRE were referred to O. subulata 
(Miihlenpfordt) Engelm. by Mr. E. J. Alexander of the New 
York Botanic Garden in 1949. Britton & Rose separate the 
two taxa, O. subulata and O. exaltata, although admitting 
they are very close. Mr. D. R. Hunt placed tropical east 
African material under O. exaltata where the South African 
material also appears to belong if the differences observed 
by Berger and Britton & Rose are of importance. 

2. Opuntia rosea DC. in Mem. Mus. 
Hist. Paris 17: 1. 15 (1828); Prodr. 3: 471 
(1828). Type: from Mexico. 

O. pallida Rose, Smithson. Misc. Coll. 50: 507 (1908). 
Type: from Mexico. 

Cylindr opuntia rosea (DC.) Backebg., Die Cactaceae 1: 
197, 1. 142 (1958). 

Plants c. 60-80 cm tall, forming much- 
branched compact bushes; the many short 
curved branches massed in upper part of plant, 
fairly equal in size. Joints cylindrical, tubercu- 
late, c. 4—5 cm in diam., densely covered with 

radiating spines (up to 20) from each raised 
areole, almost hiding the stems; spines c. 25-40 
mm long, covered with whitish sheaths. Flow- 
ers c. 4 cm long, rarely developed; hypanthium 
narrowly ovoid, the tubercles tipped with some 
minute acicular spines and claw-like leaf rudi- 
ments. Perianth segments with the inner 
petaloid, c. 10 (-15) mm long, oblong, obtuse, 
wine-red, patent to recurved with age. Stamens 
and style shortly exserted, wine red. Fruits 
often proliferous, usually sterile, obtusely 
obovoid to globose, c. 3 cm long, swollen, 
nearly smooth. 

Recorded from the Douglas district in the northern 
Cape Province, where there is a heavy infestation in dry 

Cape. — 2923 (Douglas): farm Kleinplaas, Annecke & 
Zimmermann sub PRE 42223; 39302. 

3. Opuntia imbricata (Haw.) DC. Prodr. 
3: 471 (1828); Britton & Rose, The Cactaceae 
1: 63 (1919); Phill. in Fmg. S. Afr. 15: 121, t.8 
(1940); Henderson & Anderson, Mem. Bot. 
Surv. S. Afr. 37: 223, f. 110 (1966); Lyman 
Benson in FI. Texas 2: 227 (1969). Type: 
Introduced into cultivation by Loddiges in 
1820; origin unknown. 

Cereus imbricatus Haw., Rev. PI. Succ. 70 (1821). 

Erect, branched, spreading shrubs often 
arborescent with age, up to 3 m high. Joints 
cylindrical, up to 40 cm long, 2-3 cm in diam., 
dark green with the raised prominent tubercles 
capped by areoles bearing 8-30 stellately ar- 
ranged spines, these variable in length, up to 3 
cm long, brown, minutely barbed and at first 
covered with a white papery sheath. Leaves 
subulate, 1-2 cm, early deciduous. Flowers 4-6 
cm long with the hypanthium forming an ab- 
breviated apical joint; petaloid segments red, 
rotate; stamens and style purple. Fruit obovoid, 
up to 5 cm long, yellow, tuberculate, inedible, 
long persistent; seeds pale yellow, c. 3 mm in 

Recorded as a pest from the eastern Cape, Natal, 
south-western Transvaal and north-western Cape. Distri- 
buted in the south-western States of North America and 

Transvaal. — 2725 (Bloemhof): Schweizer-Reneke, Du 
Toit sub PRE 24539. 

Cape. — 2824 (Kimberley): Kimberley Mining area, 
Leistner 3507. 3227 (Stutterheim): Cathcart, Riverside, 8 
km N.N.W. of village, Forester s.n. 

Common names: Kabelturksvy; Imbricate cactus. 



4. Opuntia aurantiaca Lindl. in Bot. 
Reg. 19, 1. 1606 (1833); Britton & Rose, The 
Cactaceae 1: 107 (1919); Lansdell in J. Dept. 
Agric. Reprint 72 (1923); Burtt Davy, FI. 
Transv. 1: 235 (1926); Phill. in Fmg. S. Afr. 13: 
216 (1938); Archibald in S. Afr. J. Sci. 36: 
195-211 (1939); Henderson & Anderson, 
Mem. Bot. Surv. S. Afr. 37: 220, f. 109 (1966). 
Type: Botanical Register, Vol. 19, t. 1606, from 
South America. 

Low spreading and creeping shrublets up 
to 1 m high, rooting where contact is made with 
the soil. Stems with a tuberous subterranean 
portion, terete above ground, much branched. 
Joints brittle, subterete to linear, 6-20 cm long, 
1-2,5 cm broad, light green; areoles close 
together, woolly, with glochidia and 2-7 sharp 
strong radiating spines, up to 3 cm long, mi- 
nutely barbed apically. Leaves rarely present. 
Flowers many, c. 4 cm long; hypanthium tubu- 
lar, usually spineless; petaloid segments lemon 
yellow to golden yellow. Fruit globose, 3-4 cm 
in diam., red, often bearing a few spines, 
inedible; able to form roots and reproduce 
vegetatively; seeds numerous, reniform, light 
brown, with low viability. Fig. 49. 

Recorded as a pest in the eastern Cape; occasional in 
the Orange Free State, Natal and Transvaal. Originally from 
South America. 

Natal. — 2830 (Dundee): Niekerkskraal, Wesselsnek, 
Paterson sub PRE 32309. 

Cape. — 3326 (Grahamstown): near Grahamstown, 

Leemann sub PRE 23290. 

According to Phillips (Fmg. in S. Afr. 13: 216, 1938) 
the species was introduced between 1850 and 1860 as an 
ornamental plant in a garden adjoining the old Mission 
Station at Hertzog, a village in the Stockenstroom district in 
the eastern Cape. (Grid 3226 DA). At present the infestation 
in the eastern Cape has become uncontrollable and eradica- 
tion by means of biological control and chemical poisons, 
while proving extremely costly, have so far met with little 
success. The taxon appears to be of hybrid origin. 

Common names: Katjie, Litjieskaktus, Litjiesturksvy, 
Platturksvy, Rankturksvy, Taaietjie, Jointed Cactus. 

5. Opuntia spinulifera Salm-Dyck, Hort. 
Dyck. 364 (1834); Britton & Rose, The Cac- 
taceae 1: 182 (1919); Phill. in Fmg. S. Afr. 15: 
120 (1940). Type: Described from plants in 
cultivation in Europe. 

Much-branched, forming large dense 
clumps up to 3 m tall (not arborescent). Joints 
flattened, orbicular, 20-40 cm in diam., 
glaucous-green, densely beset with areoles in a 
regular sub-spiral arrangment, areoles small, c. 
4 mm in diam., sunken, with numerous 

glochidia and 3-6 spines of varying lengths 
mostly pointing downwards, the longest c. 2 cm 
long, the marginal spine clusters more strongly 
developed. Flowers with a cup-shaped hypan- 
thium, 2,5 cm long wide, profusely covered by 
areoles containing bristles around the mouth; 
petaloid segments yellow, up to 4 cm long. 
Fruit not seen. (Britton & Rose do not describe 

Recorded as a pest in the eastern Cape and Natal. Also 
reported as a pest in Australia. Originally from Mexico. 

Natal. — 2930 (Pietermaritzburg): Pietermaritzburg, 

Harding 5. 

Cape. — 3226 (Fort Beaufort): Stockenstroom district, 
Oates s.n.; Hertzog and Seymour, Van der Merwe sub PRE 

Common names: Blouturksvy, Rondeblaar, Groot 
Rondeblaar, Saucepan cactus. Large Roundleaved Prickly 

6. Opuntia sp. aff. O. lindheimeri En- 

gelm. in Bost. J. Nat. Hist. 6: 207 (1850); 
Britton & Rose, The Cactaceae 1: 165 (1919); 
Phill. in Fmg. S. Afr. 15: 119, t.7 (1940); 
Lyman Benson in FI. Texas 2: 246 (1969). Type 
from Texas. 

Erect spreading shrubs 1-1,5 m tall. Joints 
flattened, orbicular to broadly obovate, c. 20 cm 
long and 16 cm broad, glaucous-green; areoles 
widely spaced, lower often atrophied, raised, c. 
5 mm in diam., woolly and with glochidia; 
spines mainly towards the apex, 1-3, up to 
about 5 cm long, about equal in length, straight 
or mostly slightly curved, stout, flattened, pale 
grey to yellow. Flowers close together above, 
in enlarged bristly areoles often attended by a 
spine; hypanthium cylindrical, with bristly and 
woolly areoles mostly around the mouth; 
sepaloid segments broadly obtriangular, apicu- 
late; petaloid segments yellow, retuse. Fruit 
obovoid, c. 7 cm long, purple, fairly smooth; 
seeds c. 5 mm in diam., white. 

Recorded as a pest in the Fort Beaufort district in the 
eastern Cape. 

Cape. — 3226 (Fort Beaufort): Seymour, Kat River Val- 
ley, Van der Merwe sub PRE 10055; Fort Beaufort, Geyer 
sub PRE 31400. 

Specimens from an infestation, confined to an area in 
the Fort Beaufort district, cannot be placed with certainty in 
any of the described species. Mr. E. J. Alexander of New 
York Botanic Gardens in 1938 identified this taxon as O. 
tardospina Griffiths. In the Flora of Texas, Lyman Benson 
reduces this species to a synonym of O. lindheimeri. This 
species is described by Britton & Rose as extremely 
variable, composed of many races, differing in armament, 
etc. This suggested that its affinity could be with this 



species, even though the spines are not described as 
flattened and curved which is the case in the local plants. 
Common names: Klein Rondeblaar, Small Roundleaved 
Prickly Pear. 

7. Opuntia dillenii ( Ker-Gawl .) Haw., 
Suppl. PI. Succ. 79 (1819); Britton & Rose, The 
Cactaceae 1: 162 (1919). Burtt Davy, FI. 
Transv. 1: 236 (1926); Phill. in Fmg. S. Afr. 15: 
119 (1940). Type: Based on an illustration by 

Cactus dillenii Ker-Gawl. in Bot. Reg. 3: t. 255 (1818). 

Low spreading bushes forming clumps or 
more upright, up to 2 m tall. Joints flattened, 
pale green, narrowly elliptic, c. 20 cm long and 
10 cm broad; areoles large, 10-12 mm in diam., 
densely fringed with bristles up to 8 mm long 
and with white curly wool in the centre; spines 
up to 8 in each areole, spreading, varying in 
length, the longest up to c. 5 cm long, straight 
or slightly curved, acicular or somewhat flat- 
tened, yellow, indistinctly banded when young, 
becoming brown with age, occasionally absent. 
Flowers with a slender curved hypanthium, 
narrowed in lower half; petaloid segments a 
dirty greenish yellow. Fruit globose, purple, c. 
5 cm long, narrowed and curved towards the 
base, with few areoles; seed c. 5 mm in diam., 

This species from Central America is now widely 
distributed through cultivation in warmer parts of the Old 
World. It is recorded as a pest in southern India and 
Australia It is fairly common around Pietermaritzburg in 
Natal, where it is known as the Pipestem Prickly Pear or 
Pypsteel-turksvy, because of the curved fruit resembling a 
tobacco pipe. 

Natal. — 2930 (Pietermaritzburg): Pentrich, near 
Pietermaritzburg, Harding 3; Nagle Dam, Paterson sub 
P RE 32671. 

Common names: Geeldoringturksvy, Pypsteelturksvy, 
Pipestem Prickly Pear. 

O. dillenii Ker-Gawl. is placed as a variety under O. 
stricta (Haw.) Haw. by Lyman Benson in Cact. Succ. J. 41: 
126 (1969), and in FI. Texas 2: 248 (1969). In South Africa 
the two taxa appear to be distinct. 

8. Opuntia stricta (Haw.) Haw., Syn. PI. 
Succ. 191 (1812); Britton & Rose, The Cac- 
taceae 1: 161 (1919); Burtt Davy, FI. Transv. 1: 
236 (1926); Lyman Benson in FI. Texas 2: 248 
(1969). Type: A cultivated plant of unknown 

Cactus strictus Haw., Misc. Nat. 188 (1803). 

Spreading much-branched dwarf bushes 
60-80 cm tall, forming thickets. Joints flat- 

tened, broadly to narrowly obovate, 8-15 cm 
long, glaucous-green, with several straight, 
spreading, acicular, yellow-brown spines up to 
4 cm long, emerging from prominent areoles, or 
spines few, small or absent. Flowers fairly 
large, c. 7 cm long, yellow, the hypanthium 
narrowly tubular, narrowed towards the base, 
straight or slightly bent, with few areoles; 
stamens and style about half as long as light 
yellow petaloid segments. Fruit narrowly 
obovoid, 4-6 cm long, purple, smooth, with 
very few or without areoles, apex hollow; seeds 
c. 5 mm diam., yellowish. 

Recorded as a pest in the eastern Cape, Natal and 
recently from northern South West Africa. It is the well 
known Pest Pear of Australia. It grows wild in western 
Cuba, Bahamas, Florida to Texas, etc. 

S.W.A. — 2016 (Otjiwarongo): Otjiwarongo, Dreyer s.n. 

Transvaal. — 2528 (Pretoria): Pretoria, wasteland, 
Codd 10760. 

Natal. — 2929 (Underberg): Estcourt, farm near town, 
Bedford s.n. 2930 (Pietermaritzburg): Pietermaritzburg, 
outskirts on road to Richmond, Hilliard sub PRE 32306. 

Cape. — 3226 (Fort Beaufort): Fort Beaufort area, Geyer 
sub PRE 32237 . 3326 (Grahamstown): around Grahams- 
town, Du Toit sub PRE 24035. 

Common Names: The Pest Pear of Australia. (Possibly 
the names Engelseturksvy, Luisies- and Suurturksvy and 
Sour Prickly Pear also apply to this species; cf. O. vul- 

9. Opuntia vulgaris Mill.,* Gard. Diet, 
ed. 8, No. 1 (1768); Britton & Rose, The 
Cactaceae 1: 156 (1919); Burtt Davy, FI. 
Transv. 1: 235 (1926); Hunt in F.T.E.A. Cac- 
taceae: 3 (1968). Type: Based on an illustration 
by Bauhin, Hist. PI. 1: 154 (1650) after L’Obel 
(Ic. 2: 241, 1591). 

Cactus opuntia L., Sp. PI. 1: 468 (1753) pro parte 
minore, quoad syn. Bauhin. 

Spreading shrubs up to 5 m tall, forming 
sturdy, often branching trunks. Joints flattened, 
usually fairly thin, bright green when young, 
variable in shape and size, narrowly obovate to 
narrowly elliptic, 10-30 cm long, 5-10 cm 
broad, attenuate below; areoles small, woolly 
with brown glochidia, spines straight, acicular, 
1-2 (-3) per areole (on the trunks up to 12), 
unequal in length, the longest 2-3 (-7) cm long, 
strong, greyish with a brown point. Flowers c. 
7-9 cm long, with a tubular, narrow hypan- 
thium attenuate at the base, spineless but with 

*Known to entomologists as O. monacantha Haw., a 
synonym of O. vulgaris. 



fairly many areoles; petaloid segments yellow densely woolly and filled with glochidia, occa- 
or the outer maroon. Fruit edible, obovoid, c. 6 sionally also bearing small spines and minute 
cm long, reddish purple with several persistent leaves; petaloid segments yellow or orange, 
areoles; seed 2 mm in diam., brown with a Fruit ellipsoid, c. 7 cm long, reddish, succulent, 
white aril. edible; seeds about 5 mm long. 

Origin unknown, probably from South America, now 
widespread. Fairly common in the Republic; planted as 
hedges around homesteads and kraals, also for fodder and 

Transvaal. — 2528 (Pretoria); Tweefontein near Pre- 
mier Mine, Du Toit sub PRE 23033. 2531 (Komatipoort): 
Komati River Poort, Van der Schijff 3994 . 

Natal. — 2632 (Bela Vista): Ndumu, near store, Stephen 
& Van Graan 1338. 2732 (Ubombo): Mkuzi Game Reserve, 
Ward 4468. 2832 (Mtubatuba): Hluhluwe Game Reserve, 
Ward 4423. 2930 (Pietermaritzburg): Pietermaritzburg, 
Harding 6. 293 1 (Stanger): Durban, Salisbury Island, Ward 
6253; Bluff, Pickworth sub PRE 32669. 

Cape. — 3325 (Port Elizabeth): Uitenhage, Petty sub PRE 
23282. 3326 (Grahamstown): Bushmans River Mouth, 
Johnston 851 ; Bathurst, Comins 2033. 

Common names: Engelseturksvy, Luisiesturksvy, 
Suurturksvy, English or Sour Prickly Pear. 

10. Opuntia ficus-indica (L.) Mill., 
Gard. Diet. ed. 8, No. 2 (1768); Britton & Rose, 
The Cactaceae 1: 177 (1919); Burtt Davy, FI. 
Transv. 1; 236 (1926); Phill. in Fmg. S. Afr. 15: 
119, t.3 (1940); Hunt in F.T.E.A. Cactaceae: 2 
(1968); Lyman Benson in FI. Texas 2: 250 
(1969). Type: Based on a plant in the Leiden 
Botanic Garden, of unknown origin. 

Cactus ficus-indicus L., Sp. PI. 1: 468 (1753). Type as 

O. maxima Mill., Gard. Diet ed. 8, No. 5 (1768); Britton 
& Rose, The Cactaceae 1: 180 (1919); Phill. in Fmg. S. Afr. 
15: 119 t.2 (1940). Type: a cultivated plant of unknown 
origin. O. megacantha Salm-Dyck, Hort. Dyck. 363 
(1834); Phill. in Fmg. S. Afr. 15: 119 (1940); Henderson & 
Anderson, Mem. Bot. Surv. S. Afr. 37: 225, fill (1966). 
Type: cultivated; probably from Mexico. 

Shrubs or trees up to 5 m tall, forming 
sturdy trunks with age. Joints flattened, nar- 
rowly elliptic to ovate, varying in size, 30-60 
cm long and 6-12 cm broad, attenuate below, 
often acute above, fairly thick, glaucous-green; 
areoles small to large, and then raised and 
woolly, with 3-6 radiating, unequally long, 
greyish white spines up to 3 (-10) cm long, 
straight or occasionally slightly curved, or 
spineless (in older plants and some cultivars). 
Leaves, if developed, minute, subulate, early 
deciduous. Flowers about 7 cm long; hypan- 
thium broadly cylindrical, contracted below, 
with numerous raised areoles spirally arranged, 

Benson believes the species to be a native of Mexico. It 
was introduced to southern Europe, Africa and India very 
long ago and is used for hedges, fodder and edible fruit. It is 
the most common and widespread species in South Africa. 

Transvaal.— 2429 (Zebediela): Sekukuniland, Magnet 
Heights, Mogg 795; Barnard sub PRE 23276; 585; 586. 
2527 (Rustenburg): Jacksonstuin near Brits, foot of 
Magaliesberg, Van Vuuren 335. 2528 (Pretoria): Pretoria, 
near Prison Reserve, De Winter s.n. 

Natal. — 2632 (Bela Vista): Ndumu Game Reserve, 
Oatley sub NH 50818. 2732 (Ubombo): Mkuzi Game 
Reserve, Ward 4450. 2930 (Pietermaritzburg): Pentrich 
Grange near Pietermaritzburg, Harding 743N; Ashburton, 
Paterson sub PRE 32307. 

O.F.S. — 2827 (Senekal): frequent on eastern slopes of 
Groot Doomkop, Goossens 935; 936. 

Cape. — 3326 (Grahamstown): Farm Belmont near 
Grahamstown, Du Toit sub PRE 23254; 23256- 23260. 

Recorded as a pest in parts of the eastern Cape, being 
especially plentiful around Uitenhage; also abundant along 
the Mediterranean and the Red Sea. Thunberg, in his Flora 
Capensis ed. Schultes on pp. XV and XVIII (1823), 
enumerates it (as Cactus Ficus) in his list of introduced 
plants he found there during his travels in the Cape, 
1772-1775, recording it as an escape, sometimes used for 

Mr W. G. Barnard in 1937-1939 described the Prickly 
Pear groves found in Sekukuniland in the eastern Transvaal 
and their importance to the native tribes living there (cf. 
notes and photos at PRE). By pruning and grazing the plants 
are encouraged to develop trunks up to 3 m high and are 
then left to branch freely, in time forming a dense canopy 
4—5 m high. At the base of the Lulu Mountains one of these 
groves, 30-60 years old, measured about 2 km by 100 m. 
“Such groves afford fine shelter against sun, wind and cold 
for farm livestock as well as pleasant play grounds for 
children. Natives depend upon the fruits for food (and beer) 
for two months in the year at the critical time between the 
current season’s grain planting and the exhaustion of the 
previous season’s stocks. Inside such groves there is a hive 
of industry — women gathering and packing fruits; girls and 
children feasting and dancing; the local chief holding an 
Indaba (Council); others milking cows and goats — in fact 
all the common activities of a tribe were being carried on 
beneath grateful shade on a scorching day in mid-summer’ ’. 
Barnard observed that old plants, 30-50 years old, became 
spineless, viz. ‘Kaalblaar’. Broken-off joints of the 
Kaalblaar, left exposed to the elements, revert to the spiny 
condition and were placed around older Kaalblaar trees to 
protect them. They are also used for fences. Curiously, a 
joint planted in good soil and well cared for may remain 
spineless. The fruits of the Kaalblaar are favoured most, 
having a better taste. He was told that, according to 
tradition, a joint was introduced into Sekukuniland from 
Kimberley in 1875. 

Common Names: Boer(e) turksvy, Kaalblaar, 

Doringblaar, Grootdoringturksvy, Mission Prickly Pear. 


Fig. 50. Pereskia aculeata, apical portion of branch showing leaves, flowers and fruits, x 1 (Pickworth 





Pereskia Mill. , Gard. Diet. Abr. ed. 4 (1754); Britton & Rose, The Cactaceae 1: 8 (1919); Vaupel 
in Pflanzenfam. ed. 2,21: 615 (1925); Hunt in Hutch., Gen. Flow. PI. 2: 433 (1967). Type species: 
P. aculeata Mill. {Cactus pereskia L.). 

Trees, shrubs or scramblers with normal stems and leaves, the raised pubescent areoles 
bearing spines in pairs or clusters. Leaves somewhat fleshy, deciduous, alternate, flat and broad. 
Flowers solitary or in terminal or axillary panicles or corymbs. Perianth rotate with the numerous 
segments entire or fringed. Stamens numerous. Ovary often with scales, spines or hairs, few- to 
many-seeded. Berry soft, globose or obovoid, unarmed or spiny; seeds black, shiny. 

Distributed in Mexico, West Indies, Central and South America, mostly in the coastal regions. A few species cultivated 
in Natal, one apparently naturalized. 

Pereskia aculeata Mill. , Gard. Diet. ed. 8 
(1768). Type: West Indies. 

Cactus pereskia L., Sp. PI. 1: 469 (1753). 

Pereskia pereskia (L.) Karsten, Deutsch. Flora 888 
(1882); Britton & Rose, The Cactaceae 1: 10 (1919). 

Shrubby to clambering with long whip-like 
branches; areoles with 1-3 straight spines near 
the base of the plant but towards the top with 
pairs of short hooked spines in the leaf axils. 
Leaves shortly petiolate, oblong to ovate, 5-7 
cm long, 3-4 cm broad, dark glossy green. 
Flowers aggregated towards the top of the 
branches, white, cream, yellow or pinkish, ro- 
tate, 2, 5-4, 5 cm in diam., with numerous 
spreading segments and stamens, strongly 
scented. Ovary beset with leafy scales, often 
spiny. Berry yellow, about 2 cm in diam., quite 

smooth when ripe; seeds black, somewhat flat- 
tened. Fig. 50. 

A garden escape in Natal and countries further north, 
forming impenetrable hedges. Planted over graves for 
protection by the Zulus. The fruits are edible and known 
under various names such as Barbados Gooseberry, Lemon 
Vine, etc. 

Transvaal. — 2229 (Waterpoort): Waterpoort area. 

Burger 109. 

Natal. — 2732 (Umbombo): Lake Sibayi, climber in 
forest, Vahrmeijer723. 2831 (Nkandla): Empangeni, fores- 
try plantation, Venter 855. 2931 (Stanger): S.E. of Stanger 
on fixed sand dunes, scrambling in a dense mass over Zulu 
graves, Smith sub PRE 32236; between Stanger and Stanger 
Beach, Ngongoni veld, fairly frequent in bush clumps, 
Acocks 10396. 3030 (Port Shepstone): Isipingo Beach, 
Ward 4926. 

Cape. — 3326 (Grahamstown): Grahamstown, near 1820 
Settler Museum, Tr ought on 100. 


Aberia Hochst 

caffra Hook.f. & Harv. . . 

edulis T. Anders 

longispina Harv 

tristis Sond 

verrucosa Hochst.* 

zeyheri Sond 

var. velutina Szyszyl. 


Acharia Thunb 

tragodes Thunb fig. 41 , 

Adenia Forsk 

buchananii Harms 

digitata (Harv.) Engl fig. 36, 

fruticosa Burn Davy 

subsp. fruticosa 

subsp. simplicifolia De Wilde 

subsp. trifoliolata De Wilde 

glauca Schinz 

gummifera (Harv.) Harms 

var. cerifera De Wilde* 

hastata (Harv.) Schinz 

var. hastata 

var. glandulifera De Wilde 

multiflora Pott 

natalensis De Wilde 

pechuelii (Engl.) Harms 

repanda (Burch.) Engl 

rhodesica Suesseng 

schlechteri Harms 

senensis (Klotzsch) Engl 

spinosa Bum Davy 

stenophylla Harms 

wilmsii Harms 

Adenogyrus Klotzsch 

krebsii Klotzsch 

Alsodeia angustifolia Thouars 

ardisiiflora Welw. ex Oliv 

ilicifolia Welw. ex Oliv 

natalensis (Engl.) Bak.f. 

Antidesma alnifolium Hook 

Aphloia (DC.) Benn 

myrtiflora Galpin 

theiformis (Vahl ) Benn fig. 26, 

subsp. madagascariensis (Clos.) Perrier var. 

closii Tul 

Augustia caffra (Meisn.) Klotzsch 

dregei (Otto & Dietr.) Klotzsch 

natalensis (Hook.) Klotzsch 

suffruticosa (Meisn.) Klotzsch 

Basananthe Peyr 

heterophylla Schinz 

littoralis Peyr. * 

pedata (Bak.f.) De Wilde fig. 38, 

polygaloides (Hutch. & Pearce) De Wilde fig. 38, 

sandersonii (Harv.) De Wilde fig. 38, 

triloba (H. Bol.) De Wilde fig. 38, 


Begonia L 

buttonii Irmscher 

caffra Meisn 


caffra sensu Burtt Davy 137 

dissecta Irmscher 141 

dregei Otto & Dietr fig. 46, 142 

var. caffra (Meisn.) A. DC 141 

var. sinuata (E. Mey. ex Otto & Dietr.) A. DC. 141 

dregei sensu Burtt Da\y 137 

favargeri Rechinger 141 

geranioides Hook.f. fig. 44, 45, 139 

gueinziana (A. DC.) Irmscher 141 

hirtella Link fig. 45, 1 39 

homonyma Steud fig. 46, 141 

natalensis Hook 142 

partita Irmscher 142 

parvifolia sensu Grah 142 

richardsiana T. Moore 142 

rudatisii Irmscher 141 

sinuata E. Mey. ex Otto & Dietr 141 

sinuata Grah 141 

sonderana Irmscher fig. 44, 45, 137 

var. transgrediens Irmscher 137 

sp 137 

suffruticosa Meisn 142 

var. gueinziana A. DC 141 

forma bolusii Irmscher 142 

forma worsdellii Irmscher 142 

sutherlandii Hook.f. fig. 44, 45, 139 

var. latior Irmscher 141 

villosa Lindl 139 

Bergia L 24 

alsinoides Holzhammer 25 

ammanioides Heyne ex Roth fig. 6, 28 

anagalloides E. Mey. ex Fenzl fig. 6, 25 

aquatica Roxb 27 

capensis L fig. 6, 27 

decumbens Planch, ex Harv fig. 6, 29 

erythroleuca Gilg 27 

glomerata Lfi fig. 6, 28 

glutinosa Dinter & Schulze-Menz fig. 6, 27 

integrifolia Dinter ex Holzhammer 25 

mossambicensis Wild 30 

palliderosa Gilg 29 

pentherana Keissl fig. 6, 29 

polyantha Sond fig. 6, 25 

prostrata Schinz 29 

salaria Brem fig. 6, 30 

sessilifolia Griseb 27 

spathulata Schinz fig. 6, 27 

verticillata Willd 27 

Blackwellia dentata Harv 74 

rufescens Am., nom. nud 75 

Blepharanthes J. E. Smith 106 

Brackenridgea A. Gray 12 

arenaria (De Wild. & Dur.) Robson fig. 3, 12 

nitida-4. Gray* 12 

Bryonia laevis Thunb 130 


Cactus dillenii Ker-Gawl 153 

ficus-indicus L 154 

opuntia L 149, 153 

pereskia L 154 

strictus Haw 153 






























































*An asterisk signifies exotic species or genera which are not naturalized; synonyms are in italics. 




Casearia Jacq 91 

gladiiformis Mast .fig. 30, 91 

junodii Schinz 91 

nitida Jacq.* 91 

Cassyta baccifera J. Mill 147 

Celastrus rotundifolius Thunb 86 

Ceratosicyos Nees 128 

ecklonii Nees 130 

laevis (Thunb.) A. Meeuse .fig. 40, 130 

Cereus imbricatus Haw 150 

martinii Labouret 145 

Chibaca salutaris Bertol. f 40 

Clemanthus Klotzsch 106 

senensis Klotzsch 114 


Cnidone mentzeloides E. Mey. ex Drege, nom. nud. 136 

Cylindropuntia rosea (DC.) Backebg 150 

Diporidium Bartl. & Wendl 1 

aboreum (Burch, ex DC.) Wendl 5 

natalitium Meisn 7 

serrulatum Hochst 9 

Doryalis E. Mey. ex Drege 84 

caffra (Hook.f. & Harv.) Warb 85 

longispina (Harv.) Warb 88 

rhamnoides (Burch, ex DC.) Warb 86 

rotundifolia (Thunb.) Warb 86 

zeyheri (Sond.) Warb 88 

Dovyalis E. Mey. ex Am. . 84 

caffra (Hook.f. & Harv.) Hook.f. . . . fig. 28, 29, 85 

celastroides sensu Sim 88 

celastroides Sond 86 

longispina (Harv.) Warb fig. 28, 29, 88 

lucida Sim fig. 29, 90 

revoluta Thom 90 

rhamnoides (Burch, ex DC.) Burch. & Harv. 

fig. 28, 29, 86 

rotundifolia (Thunb.) Thunb. & Harv. . . fig. 29, 86 

tristis (Sond.) Warb 88 

var. depauperata Sim 90 

zeyheri (Sond.) Warb fig. 29, 88 

zizyphoides E. Mey. ex Sond 

Echinothamnus Engl 106 

pechuelii Engl Ill 


Elatine L 30 

hydropiper L. * 30 

luxurious Del 27 


hirsuta sensu L 34 

krebsii Cham. & Schlechtd 34 

laevis L* 33 

nodiflora Lam 33 

nothria Thunb 34 

pomonensis Pohnert 36 

pulverulenta L 33 

repens (Berg.) Fourc fig. 8, 34 

Garcinia L 20 

angolensis Vesque 21 

ferrandii Chiov 21 

gerrardii Harv. ex Sim 21 

livingstonei T. Anders fig. 5, 21 

mangostana L 20 

natalensis Schltr. 21 

Gerrardina Oliv 72 

eylesiana Milne-Redh 74 

foliosa Oliv fig. 23, 72 

Gmelina indica Burnt, f 82 

Guthriea H. Bol 132 

capensis H. Bol fig. 42, 132 

Harrisia Britton* 145 

gracilis (Mill.) Britton* 145 

martinii (Labouret) Britton '..... fig. 47, 145 

Homalium Jacq 74 

subgenus Blackwellia Warb 74 

subgenus Homalium 74 

abdessammadii Aschers. & Schweinf. 77 

subsp. wildemanianum (Gilg) Wild 77 

chasei Wild* 75 

dentatum (Harv.) Warb 74 

macranthum Gilg 77 

racemosum Jacq. * 74 

rhodesicum Dunkley 77 

rufescens Benth fig. 24, 75 

subsuperum Sprague 74 

wildemanianum Gilg 77 

Hounea Baill 105 

Hybanthus Jacq 46 

caffer (Sond.) Engl 47 

var. angustifolius Engl 47 

capensis (Thunb.) Engl fig. 13, 49 

densifolius Engl fig. 13, 47 

enneaspermus (L.) F. Muell fig. 13, 47 

var. caffer (Sond.) Robson 47 

var. serratus Engl 47 

havanensis Jacq. * 46 

hirtus (Klotzsch) Engl 47 

triandra Schkuhr fig. 7, 32 

Erblichia Seem 100 

Eriudaphus Nees 63 

ecklonii Nees 64 

mundii Eckl. & Zeyh 65 

serratus Harv 65 

zeyheri Nees 64 

Erythrocarpus Roem 106 

Euonymus inermis Forsk 10 

Fissenia err 134 


Flacourtia L’Herit 82 

hirtiuscula Oliv 84 

indica (Burmf.) Merr fig-27, 82 

ramontchi L’Herit 82 

rhamnoides Burch, ex DC 86 


Frankenia L 33 

densa Pohnert 33 

var. glabrescens Engl 47 

natalensis (Harv.) Burtt Davy 49 

parviflorus (L.f.) Baill fig. 13, 49 

sp 49 

thorncroftii (N.E. Br.) Burtt Davy 49 

thymifolius (Presl) Engl 47 

Hypericum L 14 

aethiopicum sensu Sond 19 

aethiopicum Thunb 17 

subsp. aethiopicum fig. 4, 17 

subsp. sonderi (Bredell) Robson fig. 4, 19 

var. glaucescens Sond 19 

calycinum L. * 15 

kalmianum L. * 15 

kalmii Forsk 15 

lalandii Choisy 16 

var. lanceolata Sond 16 

var. lanceolatum Keller 16 

var. latifolia Sond 16 




var. macropetala Sond 16 

var. transvaalense Bredell 16 

lanuriense De Wild 15 

lanceolatum Lam 15 

leucoptychodes Steud. ex A. Rich 15 

monogyrum L. * 15 

natalense Wood & Evans 19 

var. petiolatum Bredell 19 

nigropunctatum Norlindh 20 

perforatum L 14 

quartinianum A. Rich. var. roeperanum (Schimp. 

ex A. Rich.) Engl 16 

revolutum Vahl 15 

roeperanum Schimp. ex A. Rich 16 

var. roeperanum 16 

var. schimperi (Hochst. ex A. Rich.) Moggi & 

Pisacchi 16 

sonderi Bredell 19 

var. transvaalense Bredell 19 

wilmsii Keller 20 

woodii Keller 19 

lonidium Vent 46 

caffrum Sond 47 

capense (Thunb.) Roem. & Schult 49 

enneaspermum (L.) Vent 47 

var. hirtum (Klotzsch) Oliv 47 

hirtum Klotzsch 47 

natalense Harv 49 

thorncroftii N.E. Br 49 

thymifolium Presl 47 

Jaggia Schinz 106 

repanda Schinz 116 

Keramanthus Hook.f. 106 

Kiggelaria L 60 

africana L fig. 18, 62 

var. obtusa (Harv.) Burtt Davy 62 

dregeana Turcz 62 

var. acuta Harv 62 

var. obtusa Harv 62 

ferruginea Eckl. & Zeyh 62 

grandifolia Warb 62 

integrifolia sensu Eckl. & Zeyh 62 

Kissenia R. Br. ex Endl 134 

capensis Endl fig. 43 , 136 

mentzeloides R. Br. ex Harv 136 

spathulata R. Br. ex T. Anders 136 

Kolbia P. Beauv 106 

Lightfootia theiformis Vahl 80 


Lundia monacantha Schumach. & Thonn 56 

Machadoa Welw. ex Benth. & Hook.f. 106 

Microblepharis (Wight & Am.) Roem 106 

Modecca Lam 106 

digitata Harv 114 

glauca Schinz, nom. nud 110 

gummifera Harv 117 

has tat a Harv 112 

paschanthus Harv 116 

repanda (Burch.) Druce 116 

senensis (Klotzsch) Mast 114 

Monospora grandifolia Hochst 78 

rotundifolia Hochst 78 

Neumannia A. Rich 80 

myrtiflora (Galpin) Th. Dur 82 

theiformis (Vahl) A. Rich 80 

Nothria repens Berg 34 



arborea Burch, ex DC 3 

var. arborea fig. 2, 5 

var. oconnorii (Phill.) Du Toit fig. 2, 5 

arenaria De Wild. & Dur 12 

aschersoniana Schinz 3 

atropurpurea sensu auct 9 

var. angustifolia Phill 9 

var. natalitia (Meisn.) Harv 7 

barbosae Robson fig. 2, 12 

chilversii Phill 7 

cinnabarina Engl. & Gilg fig. 2, 11 

confusa Burtt Davy & Greenway fig. 1,2, 7 

fuscescens Heine 3 

gamostigmata Du Toit fig. 2, 9 

glauca Verdoorn fig. 2, 10 

holstii Engl fig. 2, 5 

inermis (Forsk.) Schweinf. fig. 2, 10 

jabotapita L. * 1 

leptoclada sensu Phill 7 

multiflora sensu Williams 9 

natalitia (Meisn.) Walp fig. 1 , 2, 7 

oconnorii Phill 5 

pretoriensis Phill fig. 1 , 2, 8 

pulchra Hook fig. 1,2, 3 

rehmannii Szyszyl 3 

rogersii Hutch., nom. nud 10 

serrulata (Hochst.) Walp fig. 2, 9 

Oncoba Forsk 56 

kraussiana (Hochst.) Planch 58 

macrophylla sensu Schinz 58 

monacantha (Schumach. & Thonn.) Steud 56 

spinosa Forsk fig. 16, 56 

tettensis sensu Hook.f. ex Harv 58 

Ophiocaulon Hook.f. 106 

cissampeloides sensu Bak.f. 117 

gummifer (Harv.) Mast 117 

Opuntia Mill 149 

aurantiaca Lindl fig. 49, 152 

compressa (Salisb.) Macbr* 149 

dillenii (Ker-Gawl.) Haw 153 

exaltataH. Berger 150 

ficus-indica (L.) Mill 154 

imbricata (Haw.) DC 150 

maxima Mill 154 

megacantha Salm-Dyck 154 

pallida Rose 150 

rosea DC 150 

sp. aff. lindheimeri Engelm 152 

spinulifera Salm-Dyck 152 

stricta (Haw.) Haw 153 

subulata (Miihlenpfordt) Engelm.* 150 

tardospina Griffiths* 152 

vulgaris Mill 153 

Paropsia Noronha ex Thouars 105 

argutidens Sleumer 105 

brazzeana Bail! fig. 35, 105 

edulis Thouars* 105 

reticulata Engl 105 

var. ovatifolia Engl 105 

var. proschii Briq 105 

Paschanthus Burch 106 

jaggii Schinz 116 

repandus Burch 1 16 


Passiflora L 124 

coerulea L fig. 39, 124, 126 




edulis Sims 127 

eichlerana Mast. * 127, 128 

foetida L 126 

incamataL.* 124 

laurifolia L 124 

quadrangularis L 124, 126 

suberosa L 126 

subpeltata Ortega 127 

Pereskia Mill 156 

aculeata Mill fig. 50, 156 

pereskia (L.) Karsten 156 

Phoberos Lour 63 

ecklonii (Nees) Am. ex Presl 64 

mundii (Eckl. & Zeyh.) Presl 65 

zeyheri (Nees) Am 64 

Piriqueta Aubl 98 

capensis (Harv.) Urb fig. 33, 100 

villosa Aubl.* 98 

Prockia sect. Aphloia DC 80 

rotundifolia (Thunb.) Eckl. & Zeyh 86 

theiformis (Vahl) Willd 80 

Pseudoscolopia Gilg 70 

polyantha Gilg fig. 22, 70 

Pseudoscolopia Phill 70 

fraseri Phill 70 

Pythagorea africana E. Mey. ex Drege, nom. nud. . 75 

rufescens E. Mey. ex Am 75 

Rawsonia Harv. & Sond. 54 

lucida Harv. & Sond fig. 15, 54 

Renardia lejocarpa Turcz 78 

Rhamnicastrum Kuntze 63 

ecklonii (Nees) Kuntze 64 

mundii (Eckl. & Zeyh.) Kuntze 65 

zeyheri (Nees) Kuntze 64 

Rhipsalis Gaertn 147 

baccifera (J. Mill.) Steam fig. 48, 147 

cassutha Gaertn 147 

cassytha err. 147 

cereusculus Haw. * 147 

Rinorea Aubl 42 

angustifolia (Thouars ) Baill fig. 12, 44 

ardisiiflora (Welw. ex Oliv.) Kuntze 44 

guianensis Aubl. * 42 

ilicifolia (Welw. ex Oliv.) Kuntze fig. 12, 44 

natalensis Engl 44 

Samydaceae 53 

Schlechterina Harms 118 

mitostemmatoides Harms fig. 37, 118 

var. holzii Harms 118 

Scolopia Schreb 63 

ecklonii (Nees) Harv 64 

var. engleri (Gilg) Phill 64 

var. gerrardii (Harv.) Phill 64 

engleri Gilg 64 

flanaganii (H. Bol.) Sim fig. 21, 67 

var. oreophila Sleumer 70 

gerrardii Harv 64 

mundii (Eckl. & Zeyh.) Warb fig. 19, 65 

oreophila (Sleumer) Killick 70 

pusilla (Gaertn.) Willd.* 63 

riparia Mildbr. & Sleumer 67 

stolzii Gilg fig- 20, 67 

var. riparia (Mildbr. & Sleumer) Sleumer ... 67 

thorncroftii Phill 64 

zeyheri (Nees) Harv 64 

Streptopetalum Hochst 98 

serratum Hochst fig. 32, 98 


Tamarix L 

angolensis Niedenzu 

aphylla (L.) Karst. * 

articulata sensu Harv 

austro-africana Schinz 

engleri Arendt, ined 

gallica L* 

ramosissima Ledeb. * 

usneoides E. Mey. ex Bunge 

usneoides E. Mey. ex Drege, nom. nud. 

Trichodia Griff 

Trimeria Harv 

alnifolia (Hook.) Harv 

grandifolia (Hochst.) Warb 

rotundifolia (Hochst.) Gilg 

trinervis Harv 

Tryphostemma Harv 

arenophilum Pott 

friesii Norlindh 

harmsianum Dinter 

heterophyllum (Schinz) Engl 

longifolium Harms 

natalense Mast 

pedatum Bak.f. 

polygaloides Hutch. & Pearce 

sagittatum Hutch. & Pearce 

sandersonii Harv 

schinzianum Harms 

schlechteri Schinz 

trilobum H. Bol 

viride Hutch. & Pearce 


Turnera L 

capensis Harv 

oculata Story 

var. oculata 

var. paucipilosa Oberm 

ulmifolia L. * 


Viola L 

abyssinica Steud. ex Oliv 

capensis Thunb 

decumbens L.f. 

var. decumbens 

var. scrotiformis (DC.) Jessop . . . 

var. stipulacea Bartl 

var. tenuis Bartl 

enneasperma L 

odorata L.* 

parviflora L.f 

scrotiformis DC 

tricolor L* 

Warburgia Engl 

breyeri Pott 

salutaris (Bertol.f.) Chiov 

stuhlmannii Engl. * 

ugandensis Sprague* 

Wormskioldia Thonn 

glandulifera Klotzsch 

juttae Dinter & Urb 

lacerata Oberm 

lobata Urb 

longipedunculata Mast 

mossambicensis A. & R. Fernandes . . 
pilosa (Willd.) Schweinf. ex Urb.* . . . 



37, 39 





36, 37 

37, 39 

• fig- 9, 37 





fig- 25, 78 





















fig- 34, 102 
fig- 34, 102 



fig- 14, 50 

-fig- 14, 52 
fig- 14, 52 










fig- 10, 40 




fig. 31, 94 


fig. 31, 97 
fig- 31, 96 
fig. 31, 97 
fig. 31, 96 


Page Page 

rehmii Suesseng 96 

schinzii sensu Burtt Davy 97 

schinzii Urb .fig. 31, 96 

tenacetifolia Klotzsch .fig. 31 , 94 

Xylosma flanaganii H. Bol 67 

Xylotheca Hochst 58 

kotzei Phill 60 

kraussiana Hochst fig. 17, 58 

var. glabrifolia Wild 60 

lasiopetala Gilg 60